The effect of stand age on the accumulation of nutrients in the aboveground components of an Aleppo pine ecosystem

Nutrient dynamics of an Aleppo pine (Pinus halepensis, Mill.) ecosystem located in the Kassandra peninsula, Central Macedonia, Northern Greece, were studied using a chronosequence approach. The nutrient composition of the Aleppo pine trees, the understory evergreen broadleaves and forest floor in adjacent stands of 23, 48, 70 and over 100 years old was determined to estimate postfire nutrient losses. The concentration of nutrients in the Aleppo pine trees, except of Ca, was reduced with increasing stand age. Ca was the most abundant nutrient in the aboveground vegetation and in forest litter, followed by N, K, Mg and P. The accumulation of nutrients in the aboveground biomass was positively related to stand age. For younger stands nutrient accumulation was considerably larger in the understory vegetation as compared to the pines, due to substantial enhancement of the understory biomass and the number of understory species present. In middle-aged stands, however, nutrient accumulation in the understory and overstory vegetation reached a balance. In addition, considerable quantities of nutrients have been accumulated in the forest floor particularly in stands of 48 years old. Therefore, any destruction during the period of maximum nutrient accumulation in the forest floor will cause degradation of the ecosystem. It is postulated that the competition for nutrients between overstory and understory vegetation may be as important as competition in soil. Forest management practices leading to the direct conversion of the understory biomass into littermass would be of great significance for the sustainability of the Aleppo pine ecosystem.     

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.     

Effects of silvicultural treatments and seasonal patterns on foliar nutrients in young post-fire Pinus halepensis forest stands

Tree growth and health status appear to be related to foliar nutrient contents. Foliar nutrient concentration might be the result of a complex interaction between soil nutrients and effective availability caused by climate, water and other site and treatment effects. This study examines foliar macronutrient (N, P, K, Ca, Mg) and organic C concentrations in Aleppo pine (Pinus halepensis) needles (between 5 and 18 months old), as well as time course variability (nine dates, from July 1999 to November 2001). Variability was assessed depending on quality site (two sites, Yeste and Calasparra; SE Spain) and seven silvicultural treatments including thinning, scrubbing, pruning and particular combinations of them. Foliar macronutrient concentrations for Aleppo pine in South-eastern Spain were slightly lower (N, P, K,) or higher (Mg, Ca) than the considered as adequate ranges for Aleppo pine and Pinus genera. However, our results agree well with other normal ranges reported for Aleppo pine in Spain and for other North American Pinus species such as P. elliottii, P. taeda and P. palustris. Site, treatment and date (season) affected significantly the foliar macronutrient and C concentration, although the most important was the date, likely due to the two growth periods per year that Aleppo pine has in Mediterranean sites. Silvicultural treatments affected foliar nutrient concentration, so that the concentrations of N, P and K were higher when treatments included thinning than those that did not. The contrary was true for Mg and Ca. However, treatments did not affect the time course of the concentration, i.e., seasonality was not broken due to treatments. Moreover, the effect of the treatments was markedly high along the first year after they were applied but the differences were attenuate 2 years later. Site affected the time course of N, K, Mg and C in a different way: while for N, K and C, at the end of study period, they were higher in Calasparra than in Yeste, for Mg the contrary was true. Nutrient ratios had a different behaviour regarding to single nutrient: although date was significant, we did not appreciate seasonality. In addition, some nutrient ratios were not affected by treatments (N/P, N/K, Ca/Mg,) or by site (N/Ca, K/Ca). Average foliar N concentration and Ca/Mg ratio explained significantly the mean diameter and height growth, so that higher is the foliar N concentration and lower is Ca/Mg, higher is the growth.     

Effects of feathermoss removal, thinning and fertilization on lodgepole pine growth, soil microclimate and stand nitrogen dynamics

The influence of a continuous feathermoss layer (dominated by Pleurozium schreberi) on soil temperature, soil water potential and tree growth was assessed in a 5-year study. The feathermoss layer was manually removed from 900 m² plots in lodgepole pine (Pinus contorta) stands of northwestern Alberta. The interception and retention of nutrients by moss was estimated in N fertilization treatments with and without moss removal. As well, the potential for intraspecific competition to affect nutrient dynamics was assessed through a thinning treatment. Removal of the moss layer increased soil temperatures in summer and increased the period when soil was frost free, but the plots without moss had soil temperature as low as −13 °C in one winter period with little snow cover. Moss removal did not significantly affect N concentrations of the tree foliage but did reduce needle weight. Thinning had little effect on the rate of diameter growth after the first 3 years, but produced a significant increase in growth by year 4. Fertilization had a consistently positive effect on radial stem increments and N content of foliage, regardless of the presence of moss on the plot. Overall, the intraspecific competition between trees was apparently greater than interspecific competition between moss and trees.     

Replacement of wildfire by whole-tree harvesting in jack pine forests: Effects on soil fertility and tree nutrition

Large areas of northern coniferous forests once naturally maintained by stand-replacing wildfires have shifted to an anthropogenic disturbance regime of clearcut harvesting followed by natural or artificial regeneration, with unknown consequences for soil biogeochemical processes. We used a comparative approach to investigate the effects of whole-tree harvesting (WTH) vs. stand-replacing wildfire (WF) on soil C and nutrient availability, and nutrition and growth of the succeeding stand, in jack pine (Pinus banksiana) forests of northern Lower Michigan. We compared total carbon (C), total nitrogen (N), potential N mineralization, and extractable phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) among stands regenerated via WTH or WF in two age classes (4–7 years and 12–18 years). We also measured jack pine foliar nutrition and height growth in these same stands, as well as estimating the contribution of legacy dead wood to ecosystem nutrient capital in young stands. We found some evidence in support of our hypothesis that WTH would leave behind greater pools of soil C and N, but lower pools of P and base cations. However, the differences we observed were confined entirely to surface organic horizons, with the two disturbance regimes indistinguishable when viewed cumulatively to our maximum sampling depth of 30 cm. Estimates of nutrient pools in legacy wood inherited by young jack pine stands were also small in comparison to total soil pools (ranging from 1 to 9% depending on the element), suggesting that decomposition and nutrient release from this material is not likely to result in noticeable differences in soil fertility later in stand development. Similar levels of soil nutrients between WTH- and WF-origin stands were reflected in our measures of jack pine foliar nutrition and height growth, which were both unaffected by mode of stand origin. Results from this study suggest that soil nutrient levels following WTH fall within the natural range of variation produced by WF in these jack pine forests; however, comparison with a similar study on boreal jack pine suggests that latitudinal effects on O-horizon nutrient capital may influence the degree to which WTH matches the effects of WF on soil nutrient availability.     

Forest Fertilization Research in Finland: A Literature Review

This paper summarizes results from fertilization research carried out in Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] stands growing on mineral soil in Finland. The use of needle and soil analyses to indicate nutrient status and the need for fertilization are discussed along with the possibilities of increasing tree growth in various site types and stages of stand development. Doses, types of fertilizer, application time, growth increase, wood quality and the profitability of fertilization are also discussed. The stand response in terms of soil acidity, biological activity, soil fauna and resistance of trees to nutrient addition are all reviewed. The effects of fertilization on understorey vegetation, mushrooms and berries, and leaching loss of nutrients are also included.     

Tree species composition affects productivity and carbon dynamics of different site types in boreal forests

The objective was to analyse how differences in the initial proportions of tree species and site fertility affect carbon sequestration in living biomass and soil. We used the individual-based simulation model EFIMOD, which is able to simulate spatially explicit competition between trees for light and nutrients. Simulations were carried out for three site types with distinct initial stocks of soil nutrients. For each site, the 100-years undisturbed dynamics of monocultures and mixtures of three tree species (Betula pendula Roth, Pinus sylvestris L. and Picea abies (L.) H. Karst.) was predicted. Changes in the proportions of competing tree species were dependent on the fertility of the site: on poor sites, pine was the most competent species, while on rich sites, spruce increased its proportion during stand succession. Net primary production (NPP) and soil respiration were the highest in stands of two coniferous species and in stands with a high initial proportion of pine. Mixed stands were more productive than monocultures; the highest overyielding was observed with mixtures of two coniferous species. Simulated NPP and carbon stocks in all pools increased from poor to rich sites. The highest carbon stocks in standing biomass were observed for mixtures of conifer species and three-species mixtures; the greatest accumulation of forest floor occurred in stands with high proportions of pine.     

Retranslocation of foliar nutrients in evergreen tree species planted in a Mediterranean environment

Internal nutrient recycling through retranslocation (resorption) is important for meeting the nutrient demands of new tissue production in trees. We conducted a comparative study of nutrient retranslocation from leaves of five tree species from three genera grown in plantation forests for commercial or environmental purposes in southern Australia--Acacia mearnsii De Wild., Eucalyptus globulus Labill., E. fraxinoides H. Deane & Maiden, E. grandis W. Hill ex Maiden and Pinus radiata D. Don. Significant amounts of nitrogen, phosphorus and potassium were retranslocated during three phases of leaf life. In the first phase, retranslocation occurred from young leaves beginning 6 months after leaf initiation, even when leaves were physiologically most active. In the second phase, retranslocation occurred from mature green leaves during their second year, and in the third phase, retranslocation occurred during senescence before leaf fall. Nutrient retranslocation occurred mainly in response to new shoot production. The pattern of retranslocation was remarkably similar in the leaves of all study species (and in the phyllodes of Casuarina glauca Sieber ex Spreng.), despite their diverse genetics, leaf forms and growth rates. There was no net retranslocation of calcium in any of the species. The amounts of nutrients at the start of each pre-retranslocation phase had a strong positive relationship with the amounts subsequently retranslocated, and all species fitted a common relationship. The percentage reduction in concentration or content (retranslocation efficiency) at a particular growth phase is subject to many variables, even within a species, and is therefore not a meaningful measure of interspecific variation. It is proposed that the pattern of retranslocation and its governing factors are similar among species in the absence of interspecies competition for growth and crown structure which occurs in mixed species stands.     

Water limitations to carbon exchange in old-growth and young ponderosa pine stands

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.     

Why do large,nitrogen rich seedlings better resist stressful transplanting conditions? A physiological analysis in two functionally contrasting Mediterranean forest species

We analysed the physiological bases that explain why large and high nitrogen (N) concentration seedlings frequently have improved survival and growth relative to small seedlings in Mediterranean woodland plantations. Large seedlings of Aleppo pine (Pinus halepensis Mill.) and holm oak (Quercus ilex L.) with high N concentration (L+), and small seedlings with either high (S+) or low (S−) N concentration, were planted on two sites of different weed competition intensity that created contrasting stress conditions. Seedling survival, growth, gas exchange, N remobilization (N_R) and uptake (N_U), and water potential were assessed through the first growing season. Weeds reduced survival and growth, but seedling response to weed competition varied among phenotypes and between species. At the end of the first growing season, L+ Aleppo pine seedlings had higher survival than both small seedling types in presence of weeds but no differences were observed in absence of weeds. Mortality differences among phenotypes occurred in spring but not in summer. L+ Aleppo pines grew more than small Aleppo pines independently of weed competition. No holm oak seedling type survived in presence of weeds and no mortality differences among phenotypes where observed in absence of weeds, although L+ holm oak seedlings grew more than small seedlings. Mortality and growth differences in Aleppo pine were linked to marked physiological differences among phenotypes while physiological differences were small among holm oak phenotypes. L+ Aleppo pines had greater root growth, gas exchange, N_R, and N_U than small seedlings, irrespective of their N concentration. Seedling size in Aleppo pine had a greater role in the performance of transplanted seedlings than N concentration. The functional differences among oak phenotypes were small whereas they were large in pine seedlings, which led to smaller differences in transplanting performance in holm oak than in pine. This suggests that the nursery seedling quality improvement for planting in dry sites could depend on the species-specific phenotypic plasticity and functional strategy. Improved transplanting performance in large Aleppo pine seedlings relative to small seedlings was linked to greater gas exchange, root growth and N cycling.     

Afforestation method affects the isotopic composition of planted Pinus halepensis in a semiarid region of Spain

We used an isotopic approach to evaluate the effects of three afforestation methods on the ecophysiology of an Aleppo pine plantation in semiarid Spain. The site preparation methods tested were excavation of planting holes (H), subsoiling (S), and subsoiling with addition of urban solid refuse to soil (S + USR). Five years after plantation establishment, trees in the S + USR treatment were over three times larger than those in the S treatment, and nearly five-fold larger than those planted in holes. Differences in tree biomass per hectare were even greater due to disparities in initial planting density and pine tree mortality among treatments. Pine trees in the S + USR treatment showed higher foliar P concentration, δ¹³C and δ¹⁵N than those in the S or H treatments. Foliar δ¹⁵N data proved that trees in the S + USR treatment utilized USR as a source of nitrogen. Foliar δ¹³C and δ¹⁸O data suggest that improved nutrient status differentially stimulated photosynthesis over stomatal conductance in the pine trees of the S + USR treatment, thus enhancing water use efficiency and growth. In the spring of 2002, trees in the S + USR treatment exhibited the most negative predawn water potentials of all the treatments, indicating that the rapid early growth induced by USR accelerated the onset of intense intra-specific competition for water. The results of this study have implications for the establishment and management of Aleppo pine plantations on semiarid soils. Planting seedlings at low density and/or early thinning of pine stands are strongly recommended if fast tree growth is to be maintained beyond the first few years after USR addition to soil. Foliar C, O and N isotope measurements can provide much insight into how resource acquisition by trees is affected by afforestation techniques in pine plantations under dry climatic conditions.     

Nutrient retranslocation within the foliage of Pinus sylvestris

During the period 1983-1987, retranslocation of N, P, K, Ca, Mg, Mn, Zn, Fe, B and Al within the foliage of Scots pine (Pinus sylvestris L.) was studied in sapling, pole-stage and mature stands in eastern Finland. Needle concentrations of the mobile nutrients N, P, and K varied seasonally because of retranslocation. In unfertilized plots, needle contents of N, P and K decreased 62-92% during senescence and needle dry weight decreased 19-51%. In some years, needle contents of Ca, Mn, Zn, Fe, B and Al increased during senescence. Retranslocation was more efficient from needles with greater dry weight. In the fertilized, pole-stage stand, needle dry weight did not decrease at senescence four years after fertilization and nutrient content decreased less than in the other years. In the mature stand, there were no clear differences in needle dry weight and nutrient changes during senescence between years or between unfertilized and fertilized plots.     

Foliar nutrient retranslocation in Eucalyptus globulus

We measured patterns of change in concentrations and contents of nitrogen, phosphorus, potassium, magnesium and calcium in fully expanded leaves of young Eucalyptus globulus (Labill.) trees growing in a plantation in southeastern Australia, over a 12-month period beginning at the onset of spring. There was significant net retranslocation of mobile nutrients on a seasonal basis from green leaves, coinciding with continued growth and production of foliage. There was a close positive relationship between initial nutrient content (N, P and K) of the leaf and amount retranslocated, and a tight coupling between N and P retranslocated from leaves. Net retranslocation was significantly correlated with basal area growth increments. Artificial shading of leaves resulted in senescence and reduction in leaf mass. It also induced retranslocation of N, P and K from leaves of different ages and from different position in the canopy. Although the mechanisms underlying the effects of shading intensity on these changes were not elucidated, shading provided an experimental tool for studying retranslocation. Comparison of the results with published data for Pinus radiata (D. Don) grown in the same environment indicated a similarity between the species in patterns of change in foliar nutrient contents and in factors governing foliar nutrient retranslocation, giving rise to unifying principles.     

竹林养分循环 Ⅱ.毛竹林内降水的养分输入及其林地迳流的养分输出

在我国亚热带毛竹(Phyllostachys pubescens Mazel ex H.de Lehaie)主要产区浙、赣两省的三块试验林中对毛竹林内降水的养分输入及其小流域迳流的养分输出进行了连续四年的观察研究。结果表明,林内降水中的养分物质浓度及其输入量的季节变化明显,且与林内降水量的季节变化密切相关。林内降水中各养分物质的浓度及输入量依大小顺序列为K~+>Ca~(2+)>Mg~(2+)>NH_4~+-N>NO_3~--N>PO_4~(3-)。迳流水养分输出量的季节变化动态与迳流水量的季节增减趋势相一致,其养分物质浓度及输出量高低顺序为K~+>Mg~(2+)>速效N>PO_4~(3-)。除作为养分移动载体的林内降水及迳流水外,毛竹生长发育特性、土壤生物活动及施肥、抚育等人力措施亦对毛竹林生态系统的养分输入与输出产生影响。文中建立了浙江省庙山坞试验林林内降水中养分元素钾的年输入量GM(1,1)预测模型。     

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.     

Estimation of nutrient removals in stem-only and whole-tree harvesting of Scots pine, Norway spruce, and birch stands with generalized nutrient equations

Whole-tree harvesting (WTH), where logging residues are removed in addition to stems, is widely practised in Fennoscandian boreal forests. WTH increases the export of nutrients from forest ecosystems. The extent of nutrient removals may depend on tree species, harvesting method, and the intensity of harvesting. We developed generalized nutrient equations for Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies Karsten), and birch (Betula pendula Roth and Betula pubescens Ehrh.) stands to be able to calculate the amounts of nitrogen, phosphorus, potassium, and calcium in stems and above-ground biomass (stem and crown) as a function of stand volume. The equations were based on Fennoscandian literature data from 34 pine, 26 spruce, and 5 birch stands, and they explained, depending on the tree species and nutrient, 61–99% and 56–87% of the variation in the nutrient amounts of stems and above-ground biomass, respectively. The calculations based on the equations showed that nutrient removals caused by stem-only harvesting (SOH) and WTH per harvested stem m³ were smaller in pine than in spruce and birch stands. If the same volume of stem is harvested, nutrient removals are, in general, nearly equal at thinnings and final cuttings in SOH, but larger in thinnings than final cuttings in WTH. If the principal aim is to minimize the nutrient removals per harvested stem m³, the harvesting should be done at mature pine stands. The effect of biomass removal on overall site nutrient status depends on site-specific factors such as atmospheric deposition, weathering of minerals, and the size of the nutrient pools in the soil.     

White spruce foliar nutrient concentrations in relation to tree growth and soil nutrient amounts

Concentrations of foliar N, P, S, K, Ca, and Mg were studied in relation to stand age, tree growth, site index, and soil nutrient amounts for natural white spruce stands on a wide range of site conditions in the sub-boreal spruce zone of British Columbia, Canada. While Ca was sufficient in every sampled stand, relatively widespread deficiency in N was diagnosed. Deficiencies of other nutrients were diagnosed only on wet to very wet sites. Foliar N, P and K were negatively correlated with stand age and positively correlated with height and diameter growth. White spruce site index was positively correlated with foliar nutrients, and their relationships were quantified using a quadratic function. Foliar nutrients, except Ca, are positively correlated with soil nutrients measured in routine chemical analysis, and their relationships were quantified using Mitscherlich's function. It is recommended that the existing standards need to be modified should they be applied to nutrient diagnosis in natural white spruce stands. These standards appear too high for N and too low for P, K, and Ca.     

Biomass and nutrient allocation in Douglas-fir and amabilis fir seedlings: influence of growth rate and nutrition

Allocation of biomass and nutrients to shoots and roots was followed for three years in fast and slow growing populations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a fast growing pioneer species, and amabilis fir (Abies amabilis Dougl. ex J. Forbes), a slow growing shade-tolerant species. Seedlings were grown for three seasons in five nutrient treatments containing varying proportions of nitrogen and phosphorus (N:P). In both species, growth was greatest in the 250:20 N:P treatment followed by the 100:60 and 100:20 treatments. Vector analysis showed that, in both species, relative to the 100:20 treatment, seedlings in the 20:20 treatment were N deficient and seedlings in the 100:4 treatment were P deficient, i.e., where deficiency is defined to mean that an increase in nutrient supply increases nutrient content, nutrient concentration and plant dry weight. Seedlings in the 100:60 treatment had a higher P content than seedlings in the 100:20 treatment but the same dry weight, indicative of what Timmer and Armstrong (1987) termed luxury consumption. No nutrient retranslocation was observed in either species until the third growing season. In Douglas-fir, the greatest percentage of nutrients was exported from one-year-old shoots between May and July of the third growing season to support new growth. The total amount and percent of nutrients retranslocated was higher in Douglas-fir than in amabilis fir. Amabilis fir seedlings also exported N and P from older shoots, but this was later partially replenished. In both species, P retranslocation was greatest in treatments with a high N:P ratio. Nitrogen retranslocation was greatest in amabilis fir seedlings in treatments with a low N:P ratio, and greatest in Douglas-fir seedlings in the 250:20 and 100:60 treatments. Potassium retranslocation was correlated with seedling size. Douglas-fir retranslocated more of its shoot N reserves into new growth at the expense of older needles when soil fertility was high and sinks were strong. Otherwise, both species preferentially translocated the elements in short supply. Thus, retranslocation varied with the ecological characteristics of species, the relative availability of soil nutrients and sink strength.     

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.     

Nutrient use and uptake in Pinus taeda

We quantified nitrogen (N), phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) content, use (nutrient amount for one growth year), retranslocation (nutrients recycled before foliage senescence), uptake (use minus retranslocation), volume production per unit of uptake and fertilizer-uptake efficiency (percent applied taken up) in a 2 x 2 (nutrient and water) factorial experiment replicated four times in an 8-year-old loblolly pine (Pinus taeda L.) stand growing on a nutrient-poor sandy soil in Scotland County, North Carolina, USA. Over 14 years, we applied 1140, 168, 393, 168 and 146 kg ha(-1) of elemental N, P, K, Ca and Mg fertilizer, respectively, and an average of 710 mm year(-1) of irrigation. All plots received complete vegetation control. Fertilization about doubled tissue N, P, K and Mg contents at age 21, whereas irrigation resulted in smaller increases in nutrient contents. Maximum annual uptake was 101, 9.3, 44, 37 and 13 kg ha(-1) year(-1) and volume production per unit of nutrient uptake was 0.35, 3.5, 0.66, 1.1 and 3.1 m(3) kg(-1), for N, P, K, Ca and Mg, respectively. Irrigated plots had greater volume production per unit of N, P, K and Mg uptake than control plots, likely because irrigation allowed photosynthesis to continue during dry periods. Fertilized plus irrigated plots had less volume production per unit of these elements than the fertilized plots either because nutrient uptake exceeded the requirement for optimum growth or because available water (rainfall plus irrigation) was insufficient for the leaf area achieved with fertilization. At age 19, fertilizer-uptake efficiencies for N, P, K, Ca and Mg were 53, 24, 62, 57 and 39%, respectively, and increased with irrigation to 68, 36, 78, 116 and 55%, respectively. The scale of fertilizer uptake was likely a result of low native site nutrient availability, study longevity, measurement of all tissue components on site, a comprehensive assessment of coarse roots, and the 3-m rooting depth. Ecosystem nitrogen retention (applied nitrogen found in living plant material, litter fall and soil to 150-cm depth) was estimated at 79% at age 17, a value that would likely be greater when including soil nitrogen to rooting depth and calculating retention at age 21 when the study ended. The ecosystem retention value provides evidence that intensive site resource management can be accomplished with low likelihood of applied materials moving offsite.