Short-term effects of fertilization on photosynthesis and leaf morphology of field-grown loblolly pine following long-term exposure to elevated CO(2) concentration

We examined effects of a first nitrogen (N) fertilizer application on upper-canopy needle morphology and gas exchange in approximately 20-m-tall loblolly pine (Pinus taeda L.) exposed to elevated carbon dioxide concentration ([CO(2)]) for 9 years. Duke Forest free-air CO(2) enrichment (FACE) plots were split and half of each ring fertilized with 112 kg ha(-1) elemental N applied in two applications in March and April 2005. Measurements of needle length (L), mass per unit area (LMA), N concentration (N(l)) on a mass and an area basis, light-saturated net photosynthesis per unit leaf area (A(a)) and per unit mass (A(m)), and leaf conductance (g(L)) began after the second fertilizer application in existing 1-year-old foliage (F(O)) and later in developing current-year first-flush (F(C1)) and current-year second-flush (F(C2)) foliage. Elevated [CO(2)] increased A(a) by 43 and 52% in F(O) and F(C1) foliage, respectively, but generally had no significant effect on any other parameter. Fertilization had little or no significant effect on L, LMA, A or g(L) in F(O) foliage; although N(l) was significantly higher in fertilized trees by midsummer. In contrast, fertilization resulted in large increases in L, N(l), and A in F(C1) and F(C2) foliage, increasing A(a) by about 20%. These results suggest that, although both needle age classes accumulate N following fertilization, they use it differently-current-year foliage incorporates N into photosynthetic machinery, whereas 1-year-old foliage serves as an N store. There were no significant interaction effects of elevated [CO(2)] and fertilization on A. Elevated [CO(2)] increased the intercept of the A:N(l) relationship but did not significantly affect the slope of the relationship in either foliage age class.     

Integrating within-crown variation in net photosynthesis in loblolly and slash pine families

We examined photosynthetic characteristics of two fast- and two slow-growing half-sib families of both loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii var. elliottii Engelm.) on two sites in northern Florida to: (1) quantify variation in light-saturated net photosynthesis (Amax) associated with vertical crown position and foliage age; (2) quantify the amount and distribution of leaf area by foliage age class; and (3) determine whether photosynthetic indices, ranging from leaf-level through whole-crown Amax, were related to growth differences among species and families. In both species, leaf-level Amax was higher in more recently formed foliage both within the same year (where Amax in the third flush averaged 10 to 30% higher than Amax in the first flush) and between years (where Amax in current-year foliage averaged 20 to 40% higher than Amax in 1-year-old foliage). When expressed on a leaf area basis, Amax of current-year foliage was higher in slash pine than in loblolly pine, but Amax expressed on a mass basis did not differ between species. Loblolly pine had higher whole-tree leaf area than slash pine, whereas whole-tree Amax did not differ between species. When the mean values for fast-growing families were compared with the mean values for slow-growing families, there were no differences in leaf-level characteristics, whereas at the whole-tree level, fast-growing families had higher leaf area and whole-tree Amax than slow-growing families in both species. When comparisons were made among the individual fast- and slow-growing families, however, results were more variable. In both species, stem volume growth was strongly correlated with whole-tree Amax, with most of the strength of the correlation deriving from the relationship between volume growth and tree leaf area.     

The effects of sulfometuron on the root growth of loblolly pine

The effect of sulfometuron (Oust) on the root growth of loblolly pine (Pinus taeda L.) seedlings was studied in the field and in two soil types in the greenhouse. In the greenhouse study sulfometuron was applied to the foliage only, the soil only, and to both foliage and soil at the rates of 0.10, 0.21, and 0.42 kg ai/ha. Twenty-eight days after application root growth significantly decreased for all methods of application and with increasing rates e.g., 0.10, 0.21, and 0.42 kg ai/ha sulfometuron reduced new root length by 42%, 53%, and 64%, respectively. Application in the field at the rate of 0.30 kg ai/ha resulted in a 68% reduction in root growth. However, by the end of the growing season treated seedling root and shoot biomass were 115% and 64% greater, respectively. To ensure earlier season survival the rate of sulfometuron applied should be kept at a minimum level that will allow both weed control and adequate root growth. Low rates would be particularly important during a dry planting season.     

Changes in rates of photosynthesis and respiration during needle development of loblolly pine

Net photosynthetic rates of developing foliage and one-year-old foliage of loblolly pine (Pinus taeda L.) were measured under field conditions. In the subsequent year, net photosynthesis and dark respiration rates of current-year and one-year-old foliage were measured under controlled environmental conditions. Loblolly pine foliage grows slowly, reaching its final size 3.5 to 4 months after bud burst. Positive rates of net photosynthesis were recorded when the foliage was 13 and 18% of final length, in the controlled-environment and field study, respectively. However, because of high rates of dark respiration during the initial growth period, a positive diurnal carbon balance did not occur until foliage was about a third of final length (40 days after bud burst). Two months after bud burst, when foliage was about 55% of final length, its photosynthetic capacity exceeded that of one-year-old foliage. The highest rates of net photosynthesis were achieved when foliage was more than 90% fully expanded.     

Effects of SO2 on photosynthesis and peroxidase activity in Scots pine needles

Short-term exposure to high concentrations of SO₂ caused a temporary decrease in the photosynthetic rate, while both short-term and long-term exposure to low concentrations of SO₂ increased photo-synthesis after the exposure. Low SO₂ concentrations did not cause any change in peroxidase activity. Thus peroxidase activity does not seem a suitable method in monitoring latent injury caused by low SO₂ concentrations.     

Genetic effects on total phenolics, condensed tannins and non-structural carbohydrates in loblolly pine (Pinus taeda L.) needles

Carbon allocation to soluble phenolics (total phenolics, proanthocyanidins (PA)) and total non-structural carbohydrates (TNC; starch and soluble sugars) in needles of widely planted, highly productive loblolly pine (Pinus taeda L.) genotypes could impact stand resistance to herbivory, and biogeochemical cycling in the southeastern USA. However, genetic and growth-related effects on loblolly pine needle chemistry are not well characterized. Therefore, we investigated genetic and growth-related effects on foliar concentrations of total phenolics, PA and TNC in two different field studies. The first study contained nine different genotypes representing a range of genetic homogeneity, growing in a 2-year-old plantation on the coastal plain of North Carolina (NC), USA. The second study contained eight clones with different growth potentials planted in a 9-year-old clonal trial replicated at two sites (Georgia (GA) and South Carolina (SC), USA). In the first study (NC), we found no genetic effects on total phenolics, PA and TNC, and there was no relationship between genotype size and foliar biochemistry. In the second study, there were no differences in height growth between sites, but the SC site showed greater diameter (diameter at breast height (DBH)) and volume, most likely due to greater tree mortality (lower stocking) which reduced competition for resources and increased growth of remaining trees. We found a significant site?×?clone effect for total phenolics with lower productivity clones showing 27-30% higher total phenolic concentrations at the GA site where DBH and volume were lower. In contrast to the predictions of growth-defense theory, clone volume was positively associated with total phenolic concentrations at the higher volume SC site, and PA concentrations at the lower volume GA site. Overall, we found no evidence of a trade-off between genotype size and defense, and genetic potential for improved growth may include increased allocation to some secondary metabolites. These results imply that deployment of more productive loblolly pine genotypes will not reduce stand resistance to herbivory, but increased production of total phenolics and PA associated with higher genotype growth potential could reduce litter decomposition rates and therefore, nutrient availability.     

Atmospheric carbon dioxide, irrigation, and fertilization effects on phenolic and nitrogen concentrations in loblolly pine (Pinus taeda) needles

Concentrations of total soluble phenolics, catechin, proanthocyanidins (PA), lignin and nitrogen (N) were measured in loblolly pine (Pinus taeda L.) needles exposed to either ambient CO(2) concentration ([CO(2)]), ambient plus 175 or ambient plus 350 micromol CO(2) mol(-1) in branch chambers for 2 years. The CO(2) treatments were superimposed on a 2 x 2 factorial combination of irrigation and fertilization treatments. In addition, we compared the effects of branch chambers and open-top chambers on needle chemistry. Proanthocyanidin and N concentrations were measured in needles from branch chambers and from trees in open-top chambers exposed concurrently for two years to either ambient [CO(2)] or ambient plus 200 micromol CO(2) mol(-1) in combination with a fertilization treatment. In the branch chambers, concentrations of total soluble phenolics in needles generally increased with needle age. Concentrations of total soluble phenolics, catechin and PA in needle extracts increased about 11% in response to the elevated [CO(2)] treatments. There were no significant treatment effects on foliar lignin concentrations. Nitrogen concentrations were about 10% lower in needles from the elevated [CO(2)] treatments than in needles from the ambient [CO(2)] treatments. Soluble phenolic and PA concentrations were higher in the control and irrigated soil treatments in about half of the comparisons; otherwise, differences were not statistically significant. Needle N concentrations increased 23% in response to fertilization. Treatment effects on PA and N concentrations were similar between branch and open-top chambers, although in this part of the study N concentrations were not significantly affected by the CO(2) treatments in either the branch or open-top chambers. We conclude that elevated [CO(2)] and low N availability affected foliar chemical composition, which could in turn affect plant-pathogen interactions, decomposition rates and mineral nutrient cycling.     

Histopathology of loblolly pine needles infected by Mycosphaerella dearnessii,the brown-spot needle disease pathogen

The histopathology of green Pinus taeda needles infected with Mycosphaerella dearnessii was investigated. Cellular abnormalities observed were predominately the collapse of mesophyll cells within the limits of the symptom lesions. The epidermis, hypodermis, endodermis, transfusion tissue, and the vascular system within the lesions were not affected. Resin ducts exhibited cell necrosis and dissolution within and distal from the visible limits of lesions. An intercellular interface was present in non-collapsed mesophyll cells at lesion limits, and the lateral extension of the interface usually corresponded to the effects observed in resin ducts distal from the lesions. Non-stromatic hyphae were rare in and distal from lesion tissue, indicating the host tissues were affected in advance of the pathogen. The production of a toxin by M. dearnessii is suspected.     

马尾松不同家系光合特性的研究

对马尾松１０个家系光合特性的研究表明：（１）不同家系在叶绿素含量，叶绿素ａ／ｂ值，气孔密度，孔径，生物量等方面存在明显差异。（２）生物量和叶纱含量，气孔密度有明显的相关，两者对生物量的贡献达９５％以上；生物量和净光合速率存在一定的正相关。     

Clonal variation in crown structure, absorbed photosynthetically active radiation and growth of loblolly pine and slash pine

Crown structure, absorbed photosynthetically active radiation (APAR) and growth were analyzed in 300 replicated loblolly (Pinus taeda L.) and slash pine (Pinus elliottii Engelm. var. elliotti) clones to: (1) quantify genetic variation in crown structural traits, growth and APAR at the species, family and clonal levels; and (2) estimate within-family genetic and environmental influences on measured variables. Species and family-within-species differences were found in some growth traits, crown size, leaf area, APAR and branch angle. Loblolly pine developed larger crowns, exposed more leaf area with an acute angle, and intercepted more radiation than slash pine. Significant differences among clones within-family were found for stem volume and crown architecture. Loblolly pine and slash pine within-family, individual-tree broad-sense heritabilities ranged from 0.00 to 0.41 for growth and crown structural traits and most were between 0.10 and 0.25 when estimated from a combined analysis across families. Genetic correlations of crown size, leaf area and APAR with volume increment generally ranged from 0.60 to 0.75. This knowledge of the genetic interactions among growth and crown structural traits improves our understanding of how crown morphology affects light interception and stand development, and ultimately how these attributes can be incorporated in the selection of families or clones for the development of new crop tree ideotypes.     

Temporal and spatial patterns of net photosynthesis in 12-year-old loblolly pine five growing seasons after thinning

Physiological parameters were measured under natural light conditions and needle orientation from towers and walkways erected in the canopy of a loblolly pine (Pinus taeda L.) plantation. Four silvicultural treatments were randomly assigned to the twelve plots in the fall of 1988. Plots were thinned to a density of 731 trees per hectare or left unthinned, at a density of 2990 trees per hectare. The plots were left unfertilized or fertilized with 744 kg/ha of diammonium triple superphosphate was applied. During the fifth growing season (1993) following thinning and fertilization, needle level physiology was not different with respect to the thinning treatment for fertilized or unfertilized plots. In contrast, upper crown levels within the fertilized and unfertilized plots had significantly higher light levels and photosynthetic rates than lower crown foliage. Light levels were greater in the thinned, fertilized plots than in the unthinned, fertilized plots. In contrast, no effect of thinning on canopy light levels was found in the unfertilized plots. Within crown variation in photosynthesis was strongly dependent on canopy light levels. A strong interaction of canopy level with thinning was apparent for net photosynthesis. Loblolly pine, being a shade intolerant species, showed only small physiological differences between needles from different parts of the crown. Because of the variability found in this study, more extensive sampling is needed to correctly describe the physiology of a forest canopy with adequate precision.     

火炬松人工林间伐效应的研究

在8、12、16年生火炬松人工林中设置了不同间伐强度试验林，通过对林分树因子和间伐作业经济效益的分析表明，间伐对火炬松林分保留木个体生长具有一定的促进作用，对幼龄林的促进作用明显大于中大龄林分，随着间伐强度增大，较大径阶立木株数比例增大，但期末总收获量减少。火炬松人工林第1次间伐宜在12年生之前进行，保留1000－1100株／hm^2。在造纸材有销路的地区，初植密度较大的林分第1次间伐可在8年生时进行，保留1100－1300株／hm^2；第2次间伐的间隔期为6年，以培育大中径材为目标的大龄林分，应以弱度至中度调整为主，保留1000株／hm^2。另外，间伐有利于提高保留木材性的均质性。     

An application of mixed effects analysis to modeling thinning effects on stem profile of loblolly pine

Stem analysis data from trees in permanent sample plots were used to evaluate thinning effects on stem form using the form exponent. Thinning significantly increased the form exponent particularly in the lower bole leading to a more neiloid form. A stem profile model that accounts for changes in stem form due to thinning effects was developed. For testing hypothesis on thinning effects on stem form, direct covariance modeling with the first order autoregressive (AR(1)) specification was used to account for correlation among within-tree observations. The mixed effects analysis technique was used in stem profile modeling to indirectly account for correlation among observations. As judged by the Akaike's information criterion (AIC) and the likelihood ratio test (LRT), substantial improvements were achieved over techniques that ignore correlation thus addressing biases in the standard error of the regression parameter estimates.     

Enhanced tolerance of photosynthesis to high-light and drought stress in Pseudotsuga menziesii seedlings grown in ultraviolet-B radiation

We investigated the effects of an ambient dose of ultraviolet-B (UV-B) radiation on chamber-grown Pseudotsuga menziesii var. glauca (Beissn.) Franco (Douglas-fir) seedlings, to determine if the presence of UV-B radiation in the growth light regime induces tolerance to environmental stresses such as high light and drought. Douglas-fir seedlings were grown without UV-B radiation or with 6 kJ m-2 day-1 of biologically effective UV-B, which is ambient for the intermountain regions of Idaho. Non-stressed seedlings grown with UV-B radiation had 35% lower seedling dry mass, 36% higher concentrations of UV-B absorbing compounds per unit leaf area, 30% lower stomatal frequencies, 25% lower light-saturated photochemical efficiencies of Photosystem II and 45% lower light-saturated stomatal conductance than non-stressed seedlings grown without UV-B radiation. After 4 days of high-light stress, seedlings grown with UV-B radiation had 32% higher light-saturated carbon assimilation rates (A(CO2)) than seedlings grown without UV-B radiation. After water was withheld from the seedlings for up to 15 days, seedlings grown with UV-B radiation had 50% higher A(CO2) and 40% higher seedling water potentials than seedlings grown without UV-B radiation. The results support the hypothesis that UV-B radiation can act as an environmental signal to induce tolerance to high-light and drought stress in Douglas-fir seedlings. Possible mechanisms for the enhanced stress tolerance are discussed.     

The influence of nutrient and water availability on carbohydrate storage in loblolly pine

We quantified the effects of nutrient and water availability on monthly whole-tree carbohydrate budgets and determined allocation patterns of storage carbohydrates in loblolly pine (Pinus taeda) to test site resource impacts on internal carbon (C) storage. A factorial combination of two nutrient and two irrigation treatments were imposed on a 7-year-old loblolly pine stand in the Sandhills of North Carolina. Monthly collections of foliage, branch, stem, bark, and root tissues were made and total non-structural carbohydrate analyses were performed on samples collected in years 3 and 4 after treatment initiation. Seasonal fluxes of carbohydrates reflected the hypothesized use and storage patterns. Starch concentrations peaked in the spring in all tissues measured; however, minimum concentrations in aboveground tissue occurred in late winter while minimum concentrations in below ground tissue occurred in late fall. Increased nutrient availability generally decreased starch concentrations in current year tissue, while increasing starch in 1-year-old woody tissue. Irrigation treatments did not significantly impact carbohydrate flux. The greatest capacity for starch storage was in below ground tissue, accounting for as much as 400 kg C/ha per year, and more than 65% of the total stored starch C pool. The absolute amount of C stored as starch was significantly increased with increased nutrient availability, however, its relative contribution to the total annual C budget was not changed.     

陆地植被对紫外辐射增强的生物响应研究进展

紫外辐射（UV-B）对人类健康、水域和陆地生态系统都存在不同程度的影响。综合论述了近年来有关陆地植物对UV-B辐射响应的研究成果及进展,包括植被外部形态、光合作用和光合色素以及UV-B吸收物质等胁迫响应特征变化。另外,对植被在生态系统尺度上对UV-B辐射增强的响应以及UV-B与其它环境与生物因子间的交互效应也进行了阐述。文献分析表明,约2/3的陆地植被对UV-B辐射增强的响应表现显著,虽然存在种间和亚种间的差异,但多数的研究显示UV-B辐射增强对陆地植被生长和发展存在不利影响。最后指出如何采取措施降低UV-B辐射增强对陆地植被影响,特别是对经济作物的影响将是未来一个重要的研究课题。     

The biology of Lophodermella conjuncta Darker on Corsican pine needles

The biology of Lophodermella conjuncta on Corsican pine needles was studied. Results of a spore exclusion experiment and histological examination of needles indicated that infection of young current years needles occurred and that the fungus first colonized only the endodermis. Infection remained latent until symptoms developed, from hyphae penetrating the mesophyll from the endodermis, in July of the year following infection. Ascocarps were produced on second year needles by mid-August. Aseospores were trapped throughout the year.Infected needles were cast prematurely during the third season. The phenomenon of latent infection of pine needles by some pine needle fungi is discussed.     

火炬松、杉木混交林混交效果分析

就营造火炬松、杉木混交林(行间混交)与其纯林进行对比试验,结果表明营造混交林较营造其纯林更具优势。三种处理中,Ⅰ处理的林分蓄积量比Ⅱ处理提高43.41%,比Ⅲ处理提高122.42%;Ⅰ处理的林分总生物量比Ⅱ处理提高86.91%,比Ⅲ处理提高115.76%,为营造火炬松、杉木混交林提供可借鉴的科学理论依据。     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.     

Effects of nitrogen on leaf physiology and growth of different families of loblolly and slash pine

Leaf physiology and fractional dry weight allocation were examined in four open-pollinated families of loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Englm. var. elliottii) in response to growth under low and high N supply for six months. Nitrogen greatly influenced dry weight allocation, total mass, leaf net photosynthesis and leaf conductance in seedlings of both species. Family variation in fine root allocation was observed under low but not high N treatment, but for the majority of physiological and growth traits, family variation was stable under varying N treatment. Family rankings based on juvenile height, diameter and biomass accretion were similar to rankings based on field performance at 22 years in slash pine but not loblolly pine. Lower leaf maintenance respiration rates were associated with families exhibiting the most rapid juvenile growth.