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.     

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.     

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     

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.     

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.     

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.     

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.     

Structural variability of vascular bundles and cell wall in rattan stem

Summary Anatomical investigations on six species ofCalamus demonstrate that fibre wall thickness and fibre percentage within the vascular bundles vary more than the vascular bundle size and number per unit area. Both fibre wall thickness and fibre percentage decrease consistently from the basal (order) to the top (younger) internodes of the stem and from the periphery to the centre at a given internode. The increase of wall thickness with age is more pronounced in fibres than in cortical and ground parenchyma. The thickening of the fibre wall with increasing stem density results from the deposition of additional lamellae. This appears to impart stiffness and determines the breaking behaviour of rattan both within the stem and among the species. The fracture mode of the fibres depends on the fibrillar orientation, which differs between broad and narrow lamellae.The first author is grateful to the Director Kerala Forest Research Institute (KFRI) for approving the collaborative research programme with the Institut für Holzbiologie und Holzschutz der Bundesforschungsanstalt für Forst und Holzwirtschaft. He is also indebted to the International Development Research Centre, Canada, for sponsoring the visiting programme under the project: Rattan (India) 3-P-86-0236: KFRI/109/1987. Thanks are due to Mrs. G. Weiner for her cooperation and Mr. P.K. Thulasidas (KFRI) for technical assistance in field collection     

Production efficiency of loblolly pine and sweetgum in response to four years of intensive management

We tested the hypothesis that productivity of intensively managed loblolly pine (Pinus taeda L.) and sweetgum (Liquidambar styraciflua L.) stands is dependent not only on leaf area, but also on foliar photosynthetic rate. Effects of irrigation (irrigation treatment), irrigation with a fertilizer solution (fertigation treatment), and fertigation + pest control (loblolly pine only; fertigation + pest control treatment) on leaf physiology and growth were compared with control plots during the third and fourth growing seasons. Complete weed control was maintained on all plots. Aboveground net primary productivity of loblolly pine and sweetgum increased from 16.3 to 40.5 Mg ha(-1) and from 4.2 to 23.9 Mg ha(-1), respectively, in response to the most intensive treatment. Relative to the control treatment, neither fertigation of sweetgum nor fertigation + pest control of loblolly pine had a significant or consistent influence on foliar N concentration, quantum yield, carboxylation efficiency, net photosynthesis, stomatal conductance, or production efficiency (increment in woody biomass per unit leaf area). Irrigation increased predawn leaf water potential and photosynthesis of loblolly pine, but it had no effect on production efficiency. Leaf area was the predominant determinant of maximum productivity in these rapidly growing plantations.     

Economics of intercropping loblolly pine and switchgrass for bioenergy markets in the southeastern United States

The main objective of this study was to assess the economics of alley cropping of loblolly pine (Pinus taeda L.) and switchgrass (Panicum virgatum) in the southern United States. Assuming a price range of switchgrass between $15 and $50?Mg^?1 and yield of 12?Mg?ha^?1 year^?1, we investigated the effect of switchgrass production on the optimal forest management for loblolly pine stands under different stumpage prices. We considered the following potential scenarios: no competition between species for resources; reduced loblolly pine productivity due to competition with switchgrass; and reduced productivity of both species due to competition for nutrients, water and light. Findings also suggested that the optimal system would depend on the competitive interactions between switchgrass and loblolly pine crops, and the expected prices for each crop. Loblolly pine monoculture would be the most profitable option for landowners compared to intercropping systems with switchgrass below $30?Mg^?1. However, when switchgrass prices are ??$30?Mg^?1, landowners would be financially better off adopting intercropping if competitive interaction between crops were minimal. In order to realize higher economic returns for intercropping system, forest landowners must make some efforts in order to diminish the decline of productivity.     

Bud development and dormancy in slash and loblolly pine. II. Effects of ethephon applications

Ethephon application on four dates did not induce budset in slash and loblolly pine seedlings. Ethephon promoted slash pine budbreak by 11 days when applied in mid-November and delayed loblolly pine budbreak by 8 days when applied in late November. Mid-November and early February applications of ethephon promoted height growth of both species.     

Water relations and growth of loblolly pine seedlings planted under a shelterwood and in a clear-cut

We investigated the influence of shelterwood conditions on water relations and growth of loblolly pine (Pinus taeda L.) seedlings on two harsh sites in eastern Texas. Site I was harvested to provide four overstory density treatments (0, 2.3, 4.6 and 9.2 m(2) of residual basal area per ha). To quantify the effects of overstory competition, trenched and nontrenched subplots, each containing 25 one-year-old seedlings, were established within each overstory treatment plot, and predawn and midday water potentials (Psi(w)), seedling growth and survival were measured during the growing season. Leaf area and seedling biomass partitioning were measured at the end of the growing season. Site II was harvested to provide two overstory density treatments (0 and 6.9 m(2) ha(-1)) and planted with one-year-old loblolly pine seedlings. Seedling Psi(w), stomatal conductance (g(wv)), transpiration flux density (E), leaf area, height and survival were determined. On Site I, seedling Psi(w) increased with increasing overstory basal area, whereas trenching only substantially affected Psi(w) of seedlings in the 9.2 m(2) ha(-1) overstory treatment. Growth was not affected by overstory treatment or trenching. On Site II, Psi(w) and g(wv) were highest during the morning hours and lowest in the afternoon, whereas E peaked in the afternoon. Vapor pressure deficits and photosynthetic photon flux density were major factors in determining g(wv) differences between treatments. On individual days, the presence of an overstory increased Psi(w) and reduced both g(wv) and E. On Site II, leaf area was affected by overstory treatment throughout most of the study. We conclude that the presence of an overstory can have ameliorative effects on harsh sites at the western fringe of the loblolly pine natural range.