Effect of carbon dioxide, fertilization and irrigation on loblolly pine branch morphology

 Foliage and wood parameters of branches of 12-year-old loblolly pine (Pinus taeda L.) trees were characterized after 21 months of exposure to fertilizer, irrigation and elevated CO₂ treatments. Branches of loblolly pine trees were enclosed in plastic chambers and exposed to ambient, ambient +175 and ambient +350 umol mol^–1 CO₂ concentrations. Measurements of foliage and wood at the fascicle, flush and branch levels were made at the end of the 21 month study period. The +350 CO₂ treatment did not significantly increase fascicle radius or length but did increase the number of fascicles on the first flush. Fertilization significantly increased fascicle radius and length, while irrigation significantly increased number of fascicles and flush length of first flush. The +350 CO₂ treatment also significantly increased flush length of the first flush. Significant interaction of fertilization and irrigation with CO₂ was observed for fascicle length. Significant interactions of fertilization and irrigation were also observed for flush length, number of fascicles and fascicle length. Observed increases in fascicle radius, fascicle length, number of fascicles and flush length may have been responsible for the significantly higher flush leaf area observed for the all three treatments. Also, a combination of fertilization and irrigation increased leaf area by 82% compared to that in the control when averaged across CO₂ treatments. At the branch level +350 CO₂ treatment significantly increased shoot length but not the number of flushes on the branch. In general with the exception of bark density and total number of needle scales, neither fertilization nor irrigation had any significant effect on other branch level parameters. Results from this study indicate that with ‘global change’ an increase in CO₂ alone may increase leaf area via an increase in flush length and number of fascicles. Combining increases in CO₂ with fertilization and irrigation could greatly enhance leaf area which when coupled to observed increases in net photosynthesis as a result of elevated CO₂ could greatly increase productivity of loblolly pine trees. Received: 22 August 1996 / Accepted: 5 March 1997     

Physiological attributes of three- and four-needle fascicles of loblolly pine (<Emphasis Type="Italic">Pinus taeda</Emphasis> L.)

Key message

Fascicle types differed morphologically but had similar photosynthetic capacity on a surface area basis.

Abstract

In Pinus species, fascicles can develop with a different number of needles than what is typical. For example, Pinus taeda fascicles typically have three needles, but sometimes have two or four. Although differing fascicle morphology could be a response to changes in the environment designed to optimize carbon gain or minimize water loss, we are unaware of any work comparing physiological differences between fascicles with different numbers of needles. We compared the physiological and morphological characteristics of three- and four-needle fascicles of a loblolly pine clone with an abnormally high abundance of four-needle fascicles to better understand whether differences in needle morphology affected photosynthetic capacity or transpiration. Three- and four-needle fascicles had equal length, diameter, and volume, but four-needle fascicles had significantly greater surface area, mass, and tissue density. Equal fascicle total volume resulted in smaller per-needle volume in four-needle fascicles compared to three-needle fascicles. On a unit surface area basis, light-saturated net assimilation, stomatal conductance and transpiration were similar between the three- and four-needle fascicles although the maximum rate of carboxylation was significantly greater in four-needle fascicles. On a per-fascicle basis, four-needle fascicles had greater transpiration, stomatal conductance, and maximum rate of light-saturated net assimilation. Our results suggest that several factors, including increased tissue density and stomatal density, offset the reduction in needle volume in four-needle fascicles, resulting in similar levels of gas exchange per unit surface area in three- and four-needle fascicles.

     

Gas exchange and canopy structure of 9-year-old loblolly pine,pitch pine and pitch x loblolly hybrids

Summary Seasonal gas exchange and canopy structure were compared among 9-year-old loblolly pine (Pinus taeda L.), pitch pine (Pinus rigida Mill.), and pitch x loblolly hybrids (Pinus rigida x taeda) growing in an F2 plantation located in Critz, Va., USA. Leaf net photosynthesis, conductance, internal CO₂ concentration (c_i), water use efficiency (WUE; photosynthesis/conductance), dark respiration and the ratio of net photosynthesis/respiration did not vary among or within the three taxa. Significant differences in volume production, crown length, total crown leaf surface area and the silhouette area of shade shoots among the taxa were observed. The loblolly-South Carolina source had greater volume and crown surface area than the pitch pine, and the hybrid taxa were intermediate between the two. Although the silhouette area ratio of shade foliage varied among taxa, it was not related to volume. A strong relationship between total leaf surface area and volume was observed. Leaf conductance, c_i, WUE and leaf water potential were the physiological parameters significantly and positively correlated with volume. This study suggests that the amount of needle surface in the canopy is more important in early stand volume growth than the leaf carbon exchange rate and the degree of needle self-shading in the lower canopy.     

Phenotypic analysis of first-year traits in a pseudo-backcross {(slash x loblolly) x slash} and the open-pollinated families of the pure-species progenitors

A single test, including one pseudo-backcross (Pinus elliottii x Pinus taeda) x P. elliottii and open-pollinated families of the pure species progenitors, was established in North Central Florida in December 2007 to study the transfer of the fast-growing characteristics from a P. taeda L. (loblolly pine) parent into the P. elliottii Engelm. (slash pine) background. Several traits were measured in the first growing season: height growth, phenology, tip moth incidence, stem traits, crown architectural and needle traits. Heterosis was evaluated for each trait using analyses of variance by fitting a linear mixed model. All traits were significantly (p value < 0.05) different among families while the significance for heterosis varied by trait. Positive heterosis was found for average rate of shoot elongation (ASRE), total growth (TG), total height and number of needles per fascicle while the opposite was true for base diameter, top diameter, fascicle length, fascicle diameter, crown projected area and phenological traits (cessation, duration and day to reach 50% of the height). Average performance (i.e., no heterosis) was found for initiation of growth, number of branches, number of nodes, tip moth incidence, sheath length and specific leaf area. The analyses indicated that introgression of loblolly pine alleles into slash pine was effective and novel trait combinations were achieved. The pseudo-backcross had larger variation in early height growth than the slash pine families and was taller than all open-pollinated families at the end of the first season. Tip moth incidence was much lower than the loblolly pine family.     

SEASONAL TRENDS IN RATES OF PHOTOSYNTHESIS AND RESPIRATION OF LOBLOLLY PINE AND WHITE PINE SEEDLINGS

Mc Gregor , William H. Davis (Clemson Coll., Clemson, S.C.), and Paul J. Kramer . Seasonal trends in rates of photosynthesis and respiration of loblolly pine and white pine seedlings. Amer. Jour. Bot. 50(8): 760–765. Illus. 1963.—Seasonal trends in the rates of photosynthesis and respiration of potted loblolly and white pine seedlings were studied by measuring CO₂ exchange with an infra-red gas analyzer at intervals during a year. The seedlings were kept out-of-doors, but measurements were made indoors at 25 C and 4000 ft-c of light. Beginning in February, the rate of photosynthesis per plant of both species increased, reaching a peak in mid-September for loblolly pine and in mid-July for white pine and then declining rapidly after mid-September for both species. The spring increase began before new needles emerged and the autumn decline was not accompanied by appreciable loss of foliage. The respiration rate of loblolly pine increased steadily throughout the year. Respiration of white pine increased until mid-May and then remained fairly constant through the remainder of the year. Maximum rate of photosynthesis per unit of fascicle length occurred in July for the white pine and in May for the loblolly pine, and rates declined after September in both species. Respiration rate per unit of fascicle length showed a marked increase as spring growth began in April, then decreased to a minimum in September and increased during the winter in both species. Total chlorophyll per seedling reached a maximum in September for both species and declined slightly during winter. Total chlorophyll per unit of fascicle length increased in the spring, declined slightly in midseason, and increased again in September. Photosynthesis per unit of chlorophyll reached a maximum in May for loblolly pine and in July for white pine. Stem elongation of white pine was 88% completed by May 15. On the same date, stem elongation of loblolly pine was only 42% completed.     

Effect of shade stress on growth,morphology, and carbon dynamics of loblolly pine branches

Summary Shade treatments were applied to the terminal portions of branches in loblolly pine trees to test whether portions of branches were autonomous with respect to carbohydrates. The shade treatments reduced light by 50% and 72% compared with full sun conditions resulting in significant reductions in net photosynthesis. Branch growth (branch diameter and length, branch and needle biomass) decreased significantly within the shaded portion of the branch. Branch and needle morphology were also affected by shading. Lateral branches subtending terminal shoots were labelled with ¹⁴C 2 times during the growing season. No movement of ¹⁴C into the terminal portions of the branch was evident during the first growth flush. However, during the second flush of growth small but statistically significant amounts of ¹⁴C were imported into the terminal portion of the shaded branches from subtending laterals. It was concluded that loblolly pine shoots were usually autonomous with respect to carbohydrate supply, but that carbohydrate movement into the terminal shoot from subtending foliage could occur under conditions of very high stress.     

Variation of specific leaf area and upscaling to leaf area index in mature Scots pine

Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm² g⁻¹) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m² m⁻²) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.     

Morphological and stomatal responses of Norway spruce foliage to irradiance within a canopy depending on shoot age

Morphological and stomatal responses of Norway spruce (Picea abies) foliage to light availability were studied in respect to shoot age. Needle minor diameter (D(1), anatomical width), major diameter (D(2), anatomical thickness), dry weight (M), and tissue density index (I(D)) increased, and needle flatness (Fl) and specific leaf area (SLA) decreased with foliage age, while shade foliage demonstrated higher morphological plasticity as compared to sun foliage. Needle minor diameter, dry weight, and the ratio of total to projected leaf area increased, and needle flatness and specific leaf area decreased with daily average photosynthetic photon flux density (Q(D)). The current-year foliage exhibited the highest variation with irradiance, while the morphological plasticity decreased with needle ageing. The morphological characteristics of needles were independent of irradiance if Q(D) was above 300 μmol m(-2) s(-1). D(1) was the only linear needle characteristic which significantly changed with light availability within a canopy, and thus determined needle flatness, SLA, as well as the ratio of total to projected leaf area (TLA/PLA). Needle flatness was a characteristic responding most sensitively to the photosynthetic photon flux density, R(2) was 0.68, 0.44, and 0.49 for the current-year, 1-year-old, and 2-year-old foliage, respectively. TLA/PLA ranged from 2.2 to 4.0 depending on D(1). Variation in SLA in response to light availability can be attributed to changes both in needle shape and tissue density. Stomatal responses to photosynthetic photon flux density (Q(P)) depended on foliage type (sun or shade) and age. Sun needles demonstrated higher daily maximum leaf conductances to water vapour compared to shade needles. The shade needles responded more sensitively to changes in Q(P) at dawn and sunset than the sun needles, while older needles of both foliage types exhibited faster stomatal responses. The light-saturation of leaf conductance (g(L)) was achieved by 20 μmol m(-2) s(-1) for shade foliage, and approximately by 50 μmol m(-2) s(-1) for sun foliage. As a rule, g(L) changed in response to irradiance faster in the evening, i.e. at decreasing irradiance. Stomata were not usually completely closed in the dark before sunrise and after sunset, the phenomenon being more pronounced in older shoots and sun needles. Nightly water losses from spruce foliage are attributable primarily to older shoots, and are related to age-dependent changes in stomatal responsiveness.     

Relative contributions of crown and phenological traits to growth of a pseudo-backcross pine family ((slash?×?loblolly)?×?slash) and its pure species progenitors

One pseudo-backcross [(slash?×?loblolly)?×?slash] (BC1) and open-pollinated families of the pure species progenitors were established in a single test in North Central Florida. Multivariate analysis was used to estimate the intra-trait correlation among the taxa, and path analyses were used to determine the relative contributions of crown architectural and phenological traits to first-year height growth. The multivariate analysis indicated that BC1, slash, and loblolly pine have different relationships among the traits studied, suggesting that a separate path analysis was required for each taxon. Path analysis coefficients of determination of the final models were 0.69, 0.73, and 0.65 for the pseudo-backcross, loblolly, and slash pine families, respectively. The ranking of traits by relative magnitude of effect on total growth was, for the pseudo-backcross crown projected area (CPA), fascicle length (FL), number of nodes (NN), number of branches (NB), number of needles per fascicle (NF), and fascicle diameter. For loblolly, this was CPA, NB, FL, NN, NF, initiation, and specific leaf area. For slash, this was CPA, NN, FL, NF, and NB. The study indicated that all crown traits considered in the path analysis had moderate effects on first-year height growth, with the exceptions of the consistently large effect of CPA and the minimal effect of the phenological traits.     

Changes in leaf expansion and epidermal screening effectiveness in Liquidambar styraciflua and Pinus taeda in response to UV-B radiation

Absorption or screening of ultraviolet-B (UV-B) radiation by the epidermis may be an important protective method by which plants avoid damage upon exposure to potentially harmful UV-B radiation. In the present study we examined the relationships among epidermal screening effectiveness, concentration of UV-absorbing compounds, epidermal anatomy and growth responses in seedlings of loblolly pine (Pinus taeda L.) and sweetgum (Liquidambar styraciflua L.). Seedlings of each species were grown in a greenhouse at the University of Maryland under either no UV-B radiation or daily supplemental UV-B radiation levels of 4, 8 or 11 kJ m^?2 of biologically effective UV-B (UV-B_BE) radiation. Loblolly pine seedlings were subsequently grown in the field under either ambient or supplemental levels of UV-B radiation. At the conclusion of the growing season, measurements of epidermal UV-B screening effectiveness were made with a fiber-optic microprobe. In loblolly pine, less than 0.5% of incident UV-B radiation was transmitted through the epidermis of fascicle needles and about 1% was transmitted in primary needles. In contrast, epidermal transmittance in sweetgum ranged from about 20% in leaves not preconditioned to UV-B exposure, to about 10% in leaves grown under UV-B radiation. The concentration of UV-absorbing compounds was unaffected by UV-B exposure, but generally increased with leaf age. Increases in epidermal thickness were observed in response to UV-B treatment in loblolly pine, and this accounted for over half of the variability in UV-B screening effectiveness. In spite of the low levels of UV-B penetration into the mesophyll, delays in leaf development (both species) and final needle size (loblolly pine) were observed. Seedling biomass was reduced by supplemental UV-B radiation in loblolly pine. We hypothesize that the UV-induced growth reductions were manifested by changes in either epidermal anatomy or epidermal secondary chemistry that might negatively impact cell elongation.     

Construction cost of loblolly and ponderosa pine leaves grown with varying carbon and nitrogen availability

We grew loblolly and ponderosa pine seedlings in a factorial experiment with two CO₂ partial pressures (35 and 70 Pa), and two nitrogen treatments (1.0 and 3.5 mol m^?3 NH₄⁺), for one growing season to examine the effects of carbon and nitrogen availability on leaf construction cost. Growth in elevated CO₂ reduced leaf nitrogen concentrations by 17 to 40%, and increased C:N by 22 to 68%. Elevated N availability increased leaf N concentrations and decreased C:N. Non-structural carbohydrates increased in high-CO₂-grown loblolly seedlings, except in fascicles from low N, and in ponderosa primary and fascicle leaves grown in high N. In loblolly, increases in starch were nearly 2-fold greater than the increases in soluble sugars. In ponderosa, only the soluble sugars were affected by CO₂. Leaf construction cost (g glucose g^?1 dm) varied by 9.3% across all treatments. All of the variation in loblolly leaf construction cost could be explained by changes in non-structural carbohydrates. A model of the response of construction cost to changes in the mass of different biochemical fractions suggests that the remainder of the variation in ponderosa, not explained by non-structural carbohydrates, is probably attributable to changes in lignin, phenolic or protein concentrations.     

Effects of elevated CO2-grown loblolly pine needles on the growth, consumption, development, and pupal weight of red-headed pine sawfly larvae reared within open-topped chambers

Seedlings of loblolly pine, Pinus taeda , were grown in open-topped chambers under four levels of CO₂: two ambient and two elevated. Larvae of the red-headed pine sawfly, Neodiprion lecontei , were reared from early instar to pupation, primarily on branches within chambers. Larval growth and mortality were assessed and leaf phytochemistry samples of immature and mature leaves collected weekly. Mature leaves grown under elevated CO₂ had significant reductions in leaf nitrogen and increases in non-structural carbohydrate contents, resulting in foliage being a poorer food source for larvae, i.e. higher carbohydrate:nitrogen ratio. Nutritional constituents of immature needles were unaffected by seedling CO₂ treatment. Volatile mono- and sesquiterpenes were unrelated to plant CO₂ treatments for either leaf age class. Larval consumption of immature needles significantly increased on seedlings grown under CO₂ enrichment, while mature needle consumption was not different between the treatments. The average weight gain per larva significantly declined in late instar larvae consuming elevated CO₂-grown needles. In spite of this reduced growth, neither the days to pupation nor pupal weights were different among the CO₂ treatments. This study suggests that enriched CO₂-induced alterations in pine needle phytochemistry can affect red-headed pine sawfly performance. However, compensatory measures by larvae, such as choosing to consume more nutritious immature needles, apparently helps offset enriched CO₂-induced reductions in the leaf quality of mature needles.     

Carbon uptake and respiration in above-ground parts of a Larix decidua × leptolepis tree

Summary Shade needles of hybrid larch (Larix decidua × leptolepis) had the same rates of photosynthesis as sun needles per dry weight and nitrogen, and a similar leaf conductance under conditions of light saturation at ambient CO₂ (A_max). However, on an area basis, A_max and specific leaf weight were lower in shade than in sun needles. Stomata of sun needles limited CO₂ uptake at light saturation by about 20%, but under natural conditions of light in the shade crown, shade needles operated in a range of saturating internal CO₂ without stomatal limitation of CO₂ uptake. In both needle types, stomata responded similarly to changes in light, but shade needles were more sensitive to changes in vapor pressure deficit than sun needles. Despite a high photosynthetic capacity, the ambient light conditions reduced the mean daily (in summer) and annual carbon gain of shade needles to less than 50% of that in sun needles. In sun needles, the transpiration per carbon gain was about 220 mol mol^–1 on an annual basis. The carbon budget of branches was determined from the photosynthetic rate, the needle biomass and respiration, the latter of which was (per growth and on a carbon basis) 1.6 mol mol^–1 year^–1 in branch and stem wood. In shade branches carbon gains exceeded carbon costs (growth + respiration) by only a factor of 1.6 compared with 3.5 in sun branches. The carbon balance of sun branches was 5 times higher per needle biomass of a branch or 9 times higher on a branch length basis than shade branches. The shade foliage (including the shaded near-stem sun foliage) only contributed approximately 23% to the total annual carbon gain of the tree.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

湿地松林叶面积指数测算

对湿地松（Pinus elliotii）当年生和多年生两针一束、三针一束叶片的长度、宽度、厚度和重量分别进行量测,据此探讨不同类型叶片的叶形和比叶面积差异,并结合样地调查数据对中科院千烟洲试验站湿地松人工林的叶面积指数进行计算。结果表明：湿地松三针一束叶片合拢后横切面基本呈圆形,当年生叶和多年生叶的平均直径分别为1．688mm和1.706mm;两针一束叶片合拢后从统计学上讲横切面不是圆形,而是椭圆形,叶片厚度方向直径大于宽度方向（当年生叶厚度和宽度方向直径分别为1．580ram和1．422mm,多年生叶分别为1．568mm和1.410mm）,但如果把厚度和宽度方向直径的平均值近似成圆柱体直径计算时误差在3％以内;如果只用厚度或宽度方向直径代表平均直径计算结果会有2％。10％的误差;当年生叶和多年生叶、两针一束叶和三针一束叶之间比叶面积差别很大,计算的三种比叶面积（投影比叶面积、圆柱面比叶面积和比表面积）中,当年生叶的比叶面积明显大于多年生叶,三针一束叶片的投影比叶面积和比表面积都大于两针一束叶片,但圆柱面比叶面积恰好相反。湿地松林的叶面积指数若按投影叶面积算为3．61,按圆柱面的外表面算为5．12,按总表面积的一半算为4．52,比利用冠层分析仪测量的结果略大。     

Direct and indirect estimates of Leaf Area Index (LAI) for lodgepole and loblolly pine stands

We compared direct and indirect estimates of leaf area index (LAI) for lodgepole and loblolly pine stands. Indirect estimates of LAI using radiative methods of the LI-COR LAI-2000 Plant Canopy Analyzer (PCA) did not correlate with allometric estimates for lodgepole pine, and correlated only weakly with litter-trap estimates for loblolly pine. The PCA consistently under-estimated LAI in lodgepole pine stands with high LAI, and over-estimated LAI in the loblolly pine stands with low LAI. We developed a physical model to test the hypothesis that the PCA may under-estimate LAI in high leaf area stands because of increased foliage overlap and, therefore, increased selfshading. Radiative estimates of LAI using the PCA for the physical model were consistenly lower than allometric measures. Results from the physical model suggested that increased foliage overlap decreased the ability of the PCA to accurately estimate LAI. The relationship between allometric and radiative measures suggested an upper asymptote in LAI estimated using the PCA. The PCA may not accurately estimate LAI in stands of low or high leaf area index, and the bias or error associated with these estimates probably depends on species and canopy structure. A species specific correction factor will not necessarily correct bias in LAI estimates using the PCA.     

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

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

Growth and Nutrient Retranslocation in Needles of Radiata Pine in Relation to Nitrogen Supply

The effects of N application on tree growth and the retranslocationof N, P, and K from young needles to new growth were examinedin young radiata pine (Pinus radiata D. Don) trees. Nitrogen fertilization increased the number and size of needles,rates of shoot production, stem volume growth and tree biomass.Foliar N and P contents (µg per needle) fluctuated ina cyclic fashion with prominent phases of accumulation, retranslocationand replenishment. The patterns of these fluctuations in controland N-fertilized trees were similar, although the fluxes ofN, P and K in and out of needles were increased by N fertilization.Greater translocation (g per tree) of N and K from needles ofN fertilized trees occurred because fertilization increasedthe needle weight and the proportion of N and K retranslocatedfrom individual needles. Nitrogen fertilization increased theretranslocation of P largely as a result of higher needle mass.Trees supplied with more than adequate amounts of P in the soilretranslocated up to 58 per cent of the initial pool of P fromyoung needles. The periods of high retranslocation coincidedwith periods of high concentrations of soil mineral N and withshoot production. Conversely, the periods of rapid replenishmentof N and P into the needles coincided with the time of slowshoot growth and low concentration of soil mineral N. The growthrate of trees, rather than the availability of nutrients inthe soil was the main factor controlling retranslocation. For radiata pine, retranslocation from needles is not a mechanismspecific for coping with low soil fertility. It seems to bea mechanism which enhances the nutrient supply to apical growingpoints, especially during periods of flushing. Pinus radiata, nitrogen supply, shoot growth, nutrient fluctuations and retranslocation, nutrient use and adaptation     

Allometry and biomass of Korean pine (Pinus koraiensis) in central Korea

Aboveground tree biomass of Korean pine (Pinus koraiensis Sieb. et Zucc.) was determined for a natural forest of Korean pine and mixed deciduous trees and seven age classes of plantation forests in central Korea. Regression analyses of the dry weights of stem wood, stem bark, branches, and needles versus diameter at breast height were used to calculate regression equations of the form of log Y = a + b log X. Biomass of Korean pine in the mixed forest was 118 Mg ha(-1), and biomass in the plantations was linearly related to stand age, ranging from 52.3 Mg ha(-1) in 11 to 20-year-old stands to 317.9 Mg ha(-1) in 71 to 80-year-old stands. The proportions of stem wood and stem bark in the total aboveground biomass decreased with stand age while those of branch and needle increased. Specific leaf area of Korean pine ranging from 35.2 to 52.1 cm2 g(-1) was significantly different among crown positions and needle ages; in general, lower crown position and current needles had the greatest surface area per unit dry weight.