Herbage response to spacing of loblolly pine trees in a minimal management silvopasture in southeastern USA

Loblolly pine (Pinus taeda L.) silvopastures often are established and maintained on sites of poor soil fertility and minimal fertilizer input. Our objective was to determine whether row spacing affected yield, quality, and botanical composition of minimally managed herbage in loblolly pine early in the tree rotation. Plots were randomly located equidistant from bordering tree rows in each of eight alley width treatments that were 2.4, 3.6, 4.9, 7.3, 9.7, 12.2, 14.6 m wide, and no trees. Row spacing affected the yield, quality, and botanical composition of pasture five to six years in the rotation especially at densities exceeding 840 trees ha^–1. Botanical composition shifted from predominantly cool-season to warm-season grasses between annual first- and second-harvests, respectively, which caused seasonal differences in several yield and quality traits. Tall fescue (Festuca arundinacea Schreb.) production appeared to be unsustainable under minimal management. Herbage yield generally increased, but quality and minerals (crude protein, IVDMD, Ca, and P) tended to decrease with spacing. The 4.9 m row spacing was minimally acceptable for herbage yield and quality. System design should seek to balance tree-crop yield and quality within the context of management constraints and site productivity.This revised version was published online in November 2005 with corrections to the Cover Date.     

Effects of carbon dioxide, fertilization, and irrigation on photosynthetic capacity of loblolly pine trees

Branches of nine-year-old loblolly pine trees grown in a 2 x 2 factorial combination of fertilization and irrigation were exposed for 11 months to ambient, ambient + 175, or ambient + 350 micro mol mol(-1) CO(2). Rates of light-saturated net photosynthesis (A(max)), maximum stomatal conductance to water vapor (g(max)), and foliar nitrogen concentration (% dry mass) were assessed monthly from April 1993 until September 1993 on 1992 foliage (one-year-old) and from July 1993 to March 1994 on 1993 foliage (current-year). Rates of A(max) of foliage in the ambient + 175 CO(2) treatment and ambient + 350 were 32-47 and 83-91% greater, respectively, than that of foliage in the ambient CO(2) treatment. There was a statistically significant interaction between CO(2) treatment and fertilization or irrigation treatment on A(max) on only one measurement date for each age class of foliage. Light-saturated stomatal conductance to water vapor (g(max)) was significantly affected by CO(2) treatment on only four measurement dates. Light-saturated g(max) in winter was only 42% of summer g(max) even though soil water during winter was near field capacity and evaporative demand was low. Fertilization increased foliar N concentration by 30% over the study period when averaged across CO(2) treatments. During the study period, the ambient + 350 CO(2) treatment decreased average foliar N concentration of one-year-old foliage in the control, irrigated, fertilized and irrigated + fertilized plots by 5, 6.4, 9.6 and 11%, respectively, compared with one-year-old foliage in the corresponding ambient CO(2) treatments. The percent increase in A(max) due to CO(2) enrichment was similar in all irrigation and fertilization treatments and the effect persisted throughout the 11-month study period for both one-year-old and current-year foliage.     

Aluminum sensitivity of loblolly pine and slash pine seedlings grown in solution culture

To probe variation in Al sensitivity of two co-occurring pine species, seedlings from six full-sib families of loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Engelm.) were grown in solution culture containing 4.4 mM (high-Al) or 0.01 mM (low-Al) AlCl(3) at pH 4 for 58 days. On average, both pine species had 41% less total dry weight in the high-Al treatment than in the low-Al treatment. Stem volume growth of slash pine was more sensitive to the high-Al treatment than that of loblolly pine. In both species, the high-Al treatment inhibited root dry weight more than shoot dry weight. Within-species variation in Al sensitivity among families was greater in loblolly pine (24 to 52% inhibition of seedling dry weight) than in slash pine (35 to 47% inhibition of seedling dry weight). Foliar Al concentration was positively correlated with Al sensitivity in slash pine but not in loblolly pine; however, in both species, the concentration of Al in roots was 20-fold greater than in foliage.     

Responses of foliar gas exchange to long-term elevated CO(2) concentrations in mature loblolly pine trees

Branches of field-grown mature loblolly pine (Pinus taeda L.) trees were exposed for 2 years (1992 and 1993) to ambient or elevated CO(2) concentrations (ambient + 165 micro mol mol(-1) or ambient + 330 micro mol mol(-1) CO(2)). Exposure to elevated CO(2) concentrations enhanced rates of net photosynthesis (P(n)) by 53-111% compared to P(n) of foliage exposed to ambient CO(2). At the same CO(2) measurement concentration, the ratio of intercellular to atmospheric CO(2) concentration (C(i)/C(a)) and stomatal conductance to water vapor did not differ among foliage grown in an ambient or enriched CO(2) concentration. Analysis of the relationship between P(n) and C(i) indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase during growth in elevated CO(2) concentrations. Based on estimates derived from P(n)/C(i) curves, there were no apparent treatment differences in dark respiration, CO(2) compensation point or P(n) at the mean C(i). In 1992, foliage in the three CO(2) treatments yielded similar estimates of CO(2)-saturated P(n) (P(max)), whereas in 1993, estimates of P(max) were higher for branches grown in elevated CO(2) than in ambient CO(2). We conclude that field-grown loblolly pine trees do not exhibit downward acclimation of leaf-level photosynthesis in their long-term response to elevated CO(2) concentrations.     

Identification and quantitative analysis of stage-specific carbohydrates in loblolly pine (Pinus taeda) zygotic embryo and female gametophyte tissues

Stage-specific analyses of starch and 18 sugars, including pentoses, hexoses, disaccharides, trisaccharides, oligosaccharides and sugar alcohols, were made throughout seed development for zygotic embryo and female gametophyte (FG) tissues of loblolly pine (Pinus taeda L.). Tissue was most often analyzed in triplicate from two open-pollinated families grown in different locations and sampled in different years. Carbohydrates were analyzed by enzymatic assay, high performance liquid chromatography or gas chromatography/mass spectrometry. For all carbohydrates quantified, peak concentrations were higher in embryo tissue than in FG tissue. Significant changes in starch and sugar concentrations occurred over time, with both seed collections showing similar trends in temporal changes. Although concentrations were not always similar, embryo and FG tissues generally showed similar patterns of change in starch and sugar concentrations over time. Total starch concentration was highest during early seed development and decreased as development progressed. The major sugars contributing to osmotic potential during early seed development were D-pinitol, sucrose, fructose and glucose. During mid-seed development, D-pinitol, sucrose, fructose, glucose, melibiose and raffinose provided major contributions to the osmotic environment. During late seed development, sucrose, raffinose, melibiose, stachyose and fructose were the major contributors to osmotic potential. These data suggest stage-specific media composition for each step in the somatic embryogenesis protocol.     

Aluminum fractions in root tips of slash pine and loblolly pine families differing in Al resistance

Aluminum (Al) distribution among several cellular fractions was investigated in root tips of seedlings of one Al-resistant and one Al-sensitive family of slash pine (Pinus elliottii Engelm.) and loblolly pine (Pinus taeda L.) grown in nutrient solution containing 100 microM AlCl3 (pH 4) for 167 h. Aluminum present in 5-mm-long root tips was fractionated into cell-wall-labile (desorbed in 0.5 mM citric acid), cell-wall-bound (retained after filtering disrupted cells through 20-microm mesh) and symplasmic (filtrate following cell disruption) fractions. When averaged across both species, 12% of Al absorbed by root tips appeared in the symplasmic fraction and 88% in the apoplasmic fraction (55% as cell-wall-labile, and 33% as cell-wall-bound). On a fresh mass basis, total Al in root tips was lower in loblolly pine than in slash pine, lower in the Al-resistant slash pine family than in the Al-sensitive slash pine family, and lower in the Al-resistant families than in the Al-sensitive families across species. Although the data support the hypothesis that Al-resistant plants limit Al uptake to root apices, they do not exclude other mechanisms of Al resistance. Differential Al resistance between the species and between slash pine families may also be associated with the size of the total non-labile and cell-wall-labile Al fractions, respectively. We were unable to identify the basis for differential Al resistance in loblolly pine.     

A summary of pine straw yields and economic benefits in loblolly, longleaf and slash pine stands

Forest landowners in the southeastern United States have the opportunity to manage their loblolly, longleaf and slash pine stands for pine straw (fresh undecomposed needles; the litter layer) for non-timber revenues. Pine straw is used primarily as mulch in landscaping and has grown in revenues paid to landowners in Georgia from $15.5 million in 1999 to $81 million in 2009. Pine straw is typically sold by the acre or by the bale. Selling pine straw by the acre may be advantageous to absentee landowners. Selling pine straw by the bale can generate more annual income, but bale counts need to be accurate and bale dimensions defined. For both methods, recent (2005–2010) pine straw multi-year revenues range from $50 to $150 per acre annually. Longleaf pine straw commands the highest price per bale, followed by slash pine, and lastly loblolly pine. Per rake yields from loblolly stands tend to be 15–30?% greater than slash and longleaf pine. Pine straw raking typically starts at canopy closure continuing to the first thinning, generating from $300 per acre to over $1000 per acre in new income. This paper summarizes pine straw yields and economics in loblolly, longleaf, and slash pine stands.     

Modeling production and decay of coarse woody debris in loblolly pine plantations

Forest management can have large impacts on the production and yield of coarse woody debris (CWD) in terrestrial ecosystems, yet few modeling tools exist to inform such efforts. The goal here was to develop a set of prediction equations for use in conjunction with loblolly pine (Pinus taeda L.) modeling and inventory systems to estimate CWD yields at scales ranging from individual trees to whole plantations. Permanent field plots from a 21-year study of thinning effects on plantation growth and yield across the commercial range of the species in the southern United States were surveyed to obtain sample data on CWD volume, density, and mass. Measured CWD properties were combined with inventory records of tree mortality over the study duration to characterize CWD production, decay and yield in a series of prediction equations. The resulting equations predict CWD attributes of dead trees including dry weight (kg) and fraction of standing versus downed woody material based on the time since death (years), tree diameter at breast height (cm) and height (m) at time of death and geographic coordinates of latitude and longitude. A stand-level equation predicts total CWD yield (Mg ha⁻¹) for thinned or unthinned stands based on plantation age, stem density (trees ha⁻¹), and the average height of dominant and codominant trees (m). Piece-level equations predict dry density (kg m⁻³) or nitrogen concentration (%) of CWD pieces based on their position (standing or down), ordinal decay classes, and latitude. The tree and stand-level prediction equations are designed for use in GIS or growth and yield modeling systems. The piece-level equations are designed to be used in inventory applications that survey CWD. The equations should facilitate the accurate and facile determination of mass, carbon, and nitrogen contents of CWD in planted loblolly pine forests of the southern United States.     

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.     

Branch growth and gas exchange in 13-year-old loblolly pine (Pinus taeda) trees in response to elevated carbon dioxide concentration and fertilization

We used whole-tree, open-top chambers to expose 13-year-old loblolly pine (Pinus taeda L.) trees, growing in soil with high or low nutrient availability, to either ambient or elevated (ambient + 200 micromol mol-1) carbon dioxide concentration ([CO2]) for 28 months. Branch growth and morphology, foliar chemistry and gas exchange characteristics were measured periodically in the upper, middle and lower crown during the 2 years of exposure. Fertilization and elevated [CO2] increased branch leaf area by 38 and 13%, respectively, and the combined effects were additive. Fertilization and elevated [CO2] differentially altered needle lengths, number of fascicles and flush length such that flush density (leaf area/flush length) increased with improved nutrition but decreased in response to elevated [CO2]. These results suggest that changes in nitrogen availability and atmospheric [CO2] may alter canopy structure, resulting in greater foliage retention and deeper crowns in loblolly pine forests. Fertilization increased foliar nitrogen concentration (N(M)), but had no consistent effect on foliar leaf mass (W(A)) or light-saturated net photosynthesis (A(sat)). However, the correlation between A(sat) and leaf nitrogen per unit area (N(A) = W(A)N(M)) ranged from strong to weak depending on the time of year, possibly reflecting seasonal shifts in the form and pools of leaf nitrogen. Elevated [CO2] had no effect on W(A), N(M) or N(A), but increased A(sat) on average by 82%. Elevated [CO2] also increased photosynthetic quantum efficiency and lowered the light compensation point, but had no effect on the photosynthetic response to intercellular [CO2], hence there was no acclimation to elevated [CO2]. Daily photosynthetic photon flux density at the upper, middle and lower canopy position was 60, 54 and 33%, respectively, of full sun incident to the top of the canopy. Despite the relatively high light penetration, W(A), N(A), A(sat) and R(d) decreased with crown depth. Although growth enhancement in response to elevated [CO2] was dependent on fertilization, [CO2] by fertilization interactions and treatment by canopy position interactions generally had little effect on the physiological parameters measured.     

INFLUENCE OF FIRE ON KOREAN PINE AND SCOTS PINE TREES IN PLANTATION

The plantations of korean pine(Pinus koraensis)and scots pine(Pinus sylvesrisvar.mongolica)are mainly pure stands.Fires are gradually causing problems in theseplantations and being paid much more attention recently.Study on the influence of fire ontrees and the adaptation to fire,therefore,is of great important to probe the fire ecologicalproperties and the protection ways of these two species.The results are as follows:Bothof the species are easily damaged by fire,but korean pine is more susceptible.In the samefire,korean pine is damaged more seriously than scots pine although they have the samesize.Young individuals have low fire resistant capacity and can be damaged seriously,andolder ones have strong fire resistance and can be damaged lightly.Up-hill fire makes aserious damage to the trees distributed in up-slopes with the reason of higher fireintensity.Down-hill fire makes a serious damage to the trees distributed in down-slopcswith the reason of higher fire severity.The larger deocambium area in t     

Dry-matter partitioning in loblolly and slash pine: Effects of fertilization and irrigation

The effects of fertilization and irrigation, singly and in combination, on growth and dry-matter allocation in seedlings of Pinus taeda and P. elliottii were investigated under field conditions. Trickle irrigation was regulated to maintain soil water potential above −10 kPa and 111 g of fertilizer (N:P:K 10:10:10 plus micronutrients) were banded around each seedling on three occasions. Treatments were initiated at the beginning of the second growing-season in the field, the same year sampling was conducted. Sampling consisted of complete excavation of 15 seedlings per treatment per species beginning in early April and continued every 6 weeks through mid-November, 1986. Each seedling was measured for height, root-collar diameter, foliage, stem and root dry-matter, and total needle surface-area. Allometric growth-analysis was used to determine dry-matter partitioning among the various tree components.

The cultural treatments affected the two species differently. Loblolly pine responded to treatments by shifting dry-matter allocation from roots to shoots, with the greatest increase observed in the fertilization treatment. Slash pine showed a similar response to irrigation and to irrigation plus fertilization, but increased allocation to roots under the fertilization treatment. Allocation to stems was greater than to foliage in both species and treatments except the control loblolly pine.     

Phenological variation in height and diameter growth in provenances and families of loblolly pine

The phenology of 5- and 6-year old loblolly pine (Pinus taeda L.) trees was studied over two different growing seasons (1993 and 94) in southwest Georgia. These trees were from 7–9 open-pollinated families from each of four different provenances planted at two locations. The provenances were: Atlantic Coastal Plain (eastern SC), Gulf Hammock (north FL), Lower Gulf (south AL, MS) and Upper Gulf (north AL, MS). Provenances did not vary as to when height growth started in spring, but showed very significant differences for the date of growth cessation in fall. The Gulf Hammock source grew the most and also had the longest height growth period, while the Upper Gulf source was first to stop height growth and had the least annual height increment. Provenances were also significantly different for the date of cessation of diameter growth (a difference of 22 days between Gulf Hammock and Upper Gulf), and the order of cessation was the same as for height. Families within provenances were significantly different for date of cessation of height growth and diameter growth. When family means were considered across provenances, there was a correlation of 0.69 (p-value = 0.0001) between annual height increment and date of height growth cessation. There was a weaker association between faster growth and a longer growing season within provenances.     

Lodgepole pine: the first evidence of seed-based somatic embryogenesis and the expression of embryogenesis marker genes in shoot bud cultures of adult trees

Of the various alternatives for cloning elite conifers, somatic embryogenesis (SE) appears to be the best option. In recent years, significant areas of lodgepole pine (Pinus contorta) forest have been devastated by the mountain pine beetle (MPB) in Western Canada. In an attempt to establish an SE propagation system for MPB-resistant lodgepole pine, several families displaying varying levels of resistance were selected for experimentation involving shoot bud and immature seed explants. In bud cultures, eight embryogenic lines were induced from 2 of 15 genotypes following various treatments. Genotype had an important influence on embryogenic culture initiation, and this effect was consistent over time. These lines were identified by microscopic observation and genetic markers. Despite the abundance of early somatic embryos, the cultures have yet to develop into mature embryos. In contrast, immature zygotic embryos (ZEs) cultured from megagametophytes initiated SE at an early dominance stage via nodule-type callus in 1 of 10 genotypes. As part of the study, putative embryogenesis-specific genes, WOX2 (WUSCHELL homeobox 2) and HAP3A, were analyzed in cultures of both shoot bud explants and ZEs. On the basis of these analyses, we postulate that PcHAP3A was expressed mainly in callus and may be involved in cell division, whereas WOX2 was expressed mainly in embryonal mass (EM)-like tissues. The findings from this study, based on molecular assessment, suggest that the cell lines derived from bud cultures were truly EM. Moreover, these experimental observations suggest that PcWOX2 could be used as an early genetic marker to discriminate embryogenic cultures from callus.     

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.     

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.     

Simulating the response of mature yellow poplar and loblolly pine trees to shifts in peak ozone periods during the growing season using the TREGRO model

Multiple TREGRO simulations were conducted with meteorological data files containing different growing season peak ozone (O(3)) episodes at O(3) exposures of 1.0 and 2.0 x ambient O(3) to assess the relationship between O(3) response and the phenology of mature yellow poplar (Liriodendron tulipifera L.) and loblolly pine (Pinus taeda L.) trees. Regardless of O(3) exposure and peak O(3) episode occurrence, a peak O(3) episode in August caused the greatest reduction in carbon (C) gain in yellow poplar, whereas a peak O(3) episode in July caused the greatest reduction in C gain of loblolly pine. In both species, timing of the greatest simulated O(3) effect corresponded with the completion of the annual foliage production phenophase. Simulated C gain of yellow poplar (total tree, coarse root, and total nonstructural carbohydrate) was reduced by O(3) to a greater extent than the corresponding compartments in loblolly pine, but the opposite was true for fine roots. This differential sensitivity to O(3) reflects the fact that both C assimilation and the O(3) response of the species were parameterized according to observed field measurements of each species. The differential sensitivity to O(3) of these species may have long-term implications for species composition in southeastern USA forests.     

Simultaneous prediction of the development of loblolly pine and woody competitors in young plantations

A system for examining the effects of hardwood density and cover of herbaceous components on mean size in newly established loblolly pine plantations was developed using a site preparation study located in the piedmont province of Georgia (USA). Multispecies density models were derived to predict the sum of crown heights for eight important hardwood species using both the current or age 1 number of rootstocks and herbaceous vegetation to account for intra- and interspecific effects. The predicted sums of crown heights for the woody species were then used as predictors of mean height, diameter, and volume for loblolly pine. A seemingly unrelated regression procedure was used to compensate for correlations in error components within each system of equations that result from using predicted crown heights as regressor variables. The effects of varying sweetgum density and andropogon cover on the inter-related components of the systems based on current and age 1 competition were examined for mean height and mean diameter of the planted loblolly pines. The cumulative impacts of associated vegetation on loblolly pine growth and the relative sensitivity of height and diameter to competing vegetation were demonstrated.