Preliminary studies on establishment of genetic transformation system in loblolly pine

A transformation system was established for loblolly pine (Pinus taeda L.) mature zygotic embryos usingAgrobacterium tumefaciens. The gene coding for the β-glucuronidase (GUS) gene was introduced into loblolly pine tissues and its transient expression was detected with histochemical staining. The influences of different genotypes.Agrobacterum concentrations. and cocultivation time on GUS expression and kanamycin resistant callus and shoot regeneration were investigated. The results showed that the highest GUS expression frequency (16.3%) and shoot regeneration frequency (78%) were obtained from genotype 9–1003 withAgrobacterium concentration decreased 9 times and cocultivation time of 56 hours respectively. GUS expression was obtained in all genotypes tested. The successful expression of the GUS gene in different genotypes suggested that it will be a useful transformation system for loblolly pine. (Responsible Editor: Chai Ruihai)     

Effect of early age woody and herbaceous competition control on wood properties of loblolly pine

Early age competition control has been reported to significantly improve the growth and yield of plantation grown loblolly pine. The objective of this paper is to understand the changes in wood properties: basal area weighted whole disk SG, earlywood SG (EWSG), latewood SG (LWSG) and latewood percent (LWP) of 14 year-old trees which received early age herbaceous and hardwood competition control, using data collected from 13-sites across 4-physiographic regions in the southeastern USA. The study was laid out in a randomized complete block design and had four levels of weed control (no weed control; woody vegetation control; herbaceous vegetation control; and woody and herbaceous vegetation control), with four blocks at each site. Increment cores 12 mm in diameter were collected at breast height (1.37 m) from 9-trees in each plot and ring-by-ring SG, EWSG, LWSG and LWP measured using a X-ray densitometer. Whole disk basal area weighted SG and LWP were determined for each tree and used for analysis. A reduction in whole disk SG of 0.039 and 0.0014 and LWP of 7.38% and 3.62% was observed for trees which received total weed control compared to no weed control, for lower and upper Coastal Plain sites, respectively. For trees receiving total weed control compared to no weed control, it was observed that the diameter of the juvenile core increased by 20% on average across all physiographic regions. However, no change in the length of the juvenile period was observed among treatments other than the regional differences.     

Influence of variable organic matter retention on nutrient availability in a 10-year-old loblolly pine plantation

The effects of varying forest floor and slash retention at time of regeneration were evaluated 10 years after the establishment of a loblolly pine plantation near Millport, Alabama. Treatments included removing, leaving unaltered, or doubling the forest floor and slash material. Forest floor and litterfall mass and nutrient concentrations, available soil N, foliar nutrient concentrations and stand yield were all impacted by the treatments. Forest floor mass and nutrient contents in the doubled treatment were significantly greater than the other two treatments. The doubled treatment accumulated 25, 45 and 350% more forest floor mass and 56, 56, and 310% more N than the control treatment in the Oi, Oe, and Oa layers, respectively. The other nutrients followed similar patterns. Potentially mineralized NO₃⁻-N in the mineral soil was also significantly higher in the doubled treatment. The positive effect of doubling the forest floor on soil N availability was reflected in greater foliage production, 30% more litterfall and 25% more stand yield for this treatment. This study shows that increasing the forest floor retention has resulted in increased nutrient availability and improved tree growth.     

Emergence timing affects root-collar diameter and mortality in loblolly pine seedbeds

Eight half-sib families of loblolly pine (Pinus taeda L.) were sown in a nursery. Total germination, speed and uniformity of germination, root-collar diameters, mortality, and percent cull were recorded by family. Families which germinated quickest attained the greatest average diameters at the end of the growing season. Low variability in date of emergence led to correspondingly low variability in seedling diameter. When data for all families were combined, seedlings which emerged earliest experienced the lowest mortality, had the largest diameters, and produced the fewest culls.     

Date of sowing and emergence timing affect growth and development of loblolly pine seedlings

Loblolly pine (Pinus taeda L.) seeds were sown from February to June in Alabama, U.S.A. If sowing was conducted in May or June, seedling emergence was reduced and seedling mortality increased. Seedlings from earlier sowings grew to greater sizes, but seedlings from later sowings grew later into the fall. Within any given sowing date, the earliest seedlings to emerge grew largest. Seedlings from the February sowing formed buds in mid-summer, flushed, and in some instances flushed yet again. However, by early October, not all seedlings from May and June sowings had formed an initial bud.     

Influences of antibiotics on plantlet regeneration via organogenesis in Ioblolly pine （Pinus taeda L.）

Three antibiotics ampicillin, carbenicillin, and cefotaxime were evaluated for their effects on induction, growth, and differentiation of organogenic calli, as well as rooting of regenerated shoots of three Ioblolly pine (Pinus taeda L.) genotypes. Of the antibiotics administered, cefotaxime maximally increased the frequency of callus formation and growth rate of organogeni ccalli, carbenicillin maximally increased the frequency of shoot regeneration and the average number of adventitious shoots per piece of organogenic callus, ampicUlin maximally decreased the rooting frequency of regenerated shoots and mean number of roots per regenerated shoot, in comparison with antibiotic-free media. Compared with the control, ampicillin minimally increased the frequency of callus formation, cefotaxime minimally increased the frequency of shoot regeneration, and carbenicillin minimally decreased the rooting frequency of regenerated shoots in three Ioblolly pine genotypes tested. All three antibiotics increased the frequencies of callus formation and shoot regeneration, and reduced the rooting frequency ot regenerated shoots suggested that the establishment of an efficient Agrobacterium tumefaciens-mediated transformation protocol for stable integration of foreign genes into Ioblolly pine need to select a suitable antibiotic. This investigation could be useful for optimizing genetic transformation of conifers.     

Phosphorus fertilizer,soils, and site preparation influence loblolly pine productivity

Growth and yield of 12-year-old loblolly pine (Pinus taeda L.) plantations were compared among five subsoil textures, seven site preparation methods, with and without phosphorus fertilizer. Phosphorus broadcast before planting increased the mean size of the loblolly pine trees. However, soil differences significantly influenced stand survival, and the method of site preparation significantly affected total stand yield. Therefore, the influence of phosphorus on total stand yield depended on soil type and site preparation method. The site preparation-fertilizer combinations yielding the most volume per soil type were: on the Lucy, Ruston, and Wagram soils, shear-windrow site preparation with phosphorus; on the Kirvin and Shubuta soils, chop-burn-harrow site preparation without phosphorus; on the Calloway and Henry soils, double chopping without phosphorus; on the Sawyer soil, chopburn-harrow site preparation with phosphorus; and on the Boswell, Cadeville, Gore, and Susquehanna soils, shear-windrow-harrow site preparation without phosphorus. Soil differences and phosphorus fertilization influenced the incidence of fusiform rust in 12-year-old stands of loblolly pine.     

A biologically-consistent stand growth model for loblolly pine in the Piedmont physiographic region, USA

A theory-influenced dynamical stand growth model formulation described well the behavior of thinned and unthinned loblolly pine plantations. A simplification containing few free parameters performed as well as a fully parametrized version. It seems particularly well suited to situations where available data is scarce. The stand model can be interfaced to additional climate, nutrition and carbon cycling modules for studying the effects of a changing environment.     

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.     

Herbicides and planting date affect early performance of container-grown and bare-root loblolly pine seedlings in Alabama

The survival of bare-root and container-grown loblolly pine (Pinus taeda L.) seedlings exceeded 90% when outplanted in March at two sites in Lee County, Alabama. At both sites, soil moisture and seedling survival were greater in March than in May. A March herbicide application reduced weed biomass by 75–80% at both sites. At the moist site, herbicide application did not affect survival. However, at the drier site, a reduction in weed biomass increased both the percent soil moisture and the survival of May-planted bare-root seedlings. When soil moisture at planting time was less than 13% on a dry weight basis container-grown seedlings survived better than bare-root seedlings. At both sites, reduced weed competition resulted in greater seedling heights and diameters.Alabama Experiment Station Journal Series No. 9-85802.     

Assessing freeze damage in loblolly pine seedlings: A comparison of ethane production to electrolyte leakage

Ethane production was evaluated as a method for assessing freeze damage to loblolly pine (Pinus taeda L.) seedlings by comparing it to the widely used electrolyte leakage method. Paired measurements, first ethane production and then electrolyte leakage, were conducted on the pooled needle samples at temperatures between 0° and –12°C. Ethane production rates increased in a linear fashion with decreasing temperatures between 0° and –12°C for both Virginia Coastal Plain (R²=0.80) and Marion County, Florida (R²=0.87) seed sources. The Florida seedlings were consistently 2° to 4°C higher than the Virginia seedlings at a given ethane level. Electrolyte leakage expressed as Index of Injury initially increased with decreasing temperatures, but then leveled off at or decreased below –8°C. The log-log linear regression of ethane production against Index of Injury indicated good correspondence for both seed sources (Virginia – R²=0.81; Florida – R²=0.91). Ethane production appears more rapid and to require less sampling than does electrolyte leakage while producing comparable results to the electrolyte leakage method.     

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.     

Soil and microbial respiration in a loblolly pine plantation in response to seven years of irrigation and fertilization

Because soil CO₂ efflux or soil respiration (R_S) is the major component of forest carbon fluxes, the effects of forest management on R_S and microbial biomass carbon (C), microbial respiration (R_H), microbial activity and fine root biomass were studied over two years in a loblolly pine (Pinus taeda L.) plantation located near Aiken, SC. Stands were six-years-old at the beginning of the study and were subjected to irrigation (no irrigation versus irrigation) and fertilization (no fertilization versus fertilization) treatments since planting. Soil respiration ranged from 2 to 6 μmol m⁻² s⁻¹ and was strongly and linearly related to soil temperature. Soil moisture and C inputs to the soil (coarse woody debris and litter mass) which may influence R_H were significantly but only weakly related to R_S. No interaction effects between irrigation and fertilization were observed for R_S and microbial variables. Irrigation increased R_S, fine root mass and microbial biomass C. In contrast, fertilization increased R_H, microbial biomass C and microbial activity but reduced fine root biomass and had no influence on R_S. Predicted annual soil C efflux ranged from 8.8 to 10.7 Mg C ha⁻¹ year⁻¹ and was lower than net primary productivity (NPP) in all stands except the non-fertilized treatment. The influence of forest management on R_S was small or insignificant relative to biomass accumulation suggesting that NPP controls the transition between a carbon source and sink in rapidly growing pine systems.     

Thinning method and intensity influence long-term mortality trends in a red pine forest

Tree mortality shapes forest development, but rising mortality can represent lost production or an adverse response to changing environmental conditions. Thinning represents a strategy for reducing mortality rates, but different thinning techniques and intensities could have varying impacts depending on how they alter stand structure. We analyzed trends in stand structure, relative density, stand-scale mortality, climate, and correlations between mortality and climate over 46 years of thinning treatments in a red pine forest in Northern Minnesota, USA to examine how thinning techniques that remove trees of different crown classes interact with growing stock manipulation to impact patterns of tree mortality. Relative density in unharvested plots increased during the first 25 years of the study to around 80%, then began to plateau, but was lower (12–62%) in thinned stands. Mortality in unharvested plots claimed 2.5 times more stems yr⁻¹ and 8.6 times as large a proportion of annual biomass increment during the last 21 years of the study compared to the first 25 years, but showed few temporal trends in thinned stands. Mortality in thinning treatments was generally lower than in controls, particularly during the last 21 years of the study when mortality averaged about 0.1% of stems yr⁻¹ and 4% of biomass increment across thinning treatments, but 0.8% of stems yr⁻¹ and 49% of biomass increment in unharvested plots. Treatments that combined thinning from above with low growing stock levels represented an exception, where mortality exceeded biomass production after initial thinning. Mortality averaged less than 0.1% of stems yr⁻¹ and less than 1% of annual biomass production in stands thinned from below. These trends suggest thinning from below minimizes mortality across a wide range of growing stock levels while thinning from above to low growing stock levels can result in dramatic short-term increases in mortality. Moderate to high growing stock levels (21–34 m² ha⁻¹) may offer greater flexibility for limiting mortality across a range of thinning methods. Mean and maximum annual and growing season temperatures rose by 0.6–1.8 °C during the study, and temperature variables were positively correlated with mortality in unharvested plots. Mortality increases in unharvested plots, however, were consistent with self-thinning principles and probably not driven by rising temperatures. These results suggest interactions between thinning method and intensity influence mortality reductions associated with thinning, and demonstrate the need for broader consideration of developmental processes as potential explanations for increased tree mortality rates in recent decades.     

Southern pine beetle regional outbreaks modeled on landscape, climate and infestation history

The southern pine beetle (Dendroctonus frontalis, SPB) is the major insect pest of pine species in the southeastern United States. It attains outbreak population levels sufficient to mass attack host pines across the landscape at scales ranging from a single forest stand to interstate epidemics. This county level analysis selected and examined the best climatic and landscape variables for predicting infestations at regional scales. The analysis showed that, for a given county, the most important factor in predicting outbreaks was that the county was classified as in outbreak status in the previous year. Other important factors included minimum winter temperature and the greatest difference between the average of daily minimums and a subsequent low temperature point, precipitation history either seasonally in the previous year or difference from average over the previous 2 years, the synchronizing effect of seasonal temperatures on beetle populations and the relative percentage of total forest area composed of host species. The statistical models showed that climatic variables are stronger indicators of outbreak likelihood than landscape structure and cover variables. Average climatic conditions were more likely to lead to outbreaks than extreme conditions, supporting the notion of coupling between a native insect and its native host. Still, some extreme events (i.e., periods of very low temperature or very high precipitation) did precede beetle infestation. This analysis suggested that there are predisposing and inciting factors at the large scale but the driving factors leading to individual infestations operate at smaller scales.     

Effect of handling and water stress on water status and rooting of loblolly pine stem cuttings

Two experiments were conducted to determine theeffect of handling, short-term storage, andinitial water stress on cutting water potential (_W) and rooting of loblolly pine(Pinus taeda L.) stem cuttings. First,stock plants and cuttings were measured for_W at predawn (04:00 a.m.) and earlymorning (09:00 a.m.). Cuttings were thensevered, wrapped in wet paper towels, andplaced in insulated containers for 2 or 7 h atapproximately 30 °C or for 21 h in coldstorage (4 °C). Water potentials ofcuttings were measured at the end of eachstorage period. Second, effects of initialwater stress on rooting performance of cuttingswere tested by withholding water from dormant(winter) and succulent (summer) cuttings forvarying periods of time. After each dryingtreatment, _W was measured on asample of cuttings and the remainder of thecuttings were transferred to a greenhouse withintermittent mist for 12 weeks.Storage of cuttings for long periods (7 to 21h) of time under low vapor pressure deficitconditions resulted in less negative waterpotentials of the cuttings. Dormant cuttingsrooted at higher percentages, even after beingexposed to lower values of _W Thelower values of _W in dormantcuttings could be attributed to higher ambientvapor pressure deficit during the drying phase. Results suggest that subjecting cuttings tomoderate water stress for a short period oftime does not adversely affect the rooting ofcuttings. Cutting water potentials below –1.7MPa appeared to reduce rooting of succulentcuttings and water potentials below –2.0 MPaaffected rooting in dormant cuttings.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

Potential effects of climate change on the growth of lodgepole pine across diameter size classes and ecological regions

We examined 65 lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) sites in Alberta using a dendrochronological approach in order to examine the relationships between climate and growth of lodgepole pine across elevational ecoregions and diameter size classes. The 4 elevational ecoregions sampled included the Boreal Highlands (BH: 13 sites); the Foothills (FH: 36 sites); a grouping of the montane and subalpine zones of the southern Rocky Mountains (RM: 12 sites); and the montane zone of the Cypress Hills (CH: 4 sites). The first diameter size class was comprised of the three largest (top) diameter trees at each site. The tree list of each plot was ranked, then divided into three groups of equal basal area and the tree at the midpoint of each group (small, medium, and large) was selected for the other three size classes. Annual growth in basal area of lodgepole pine was generally sensitive to heat and moisture stress in late summer of the previous year, the degree of winter harshness, and the timing of the start of the growing season. Growth was inhibited by low temperature in all winter months at the most northern BH sites which had the coldest winters, but this effect was interrupted in some of the midwinter months in the more southerly sites in the RM, and we postulated this was due to the damaging influences of Chinook winds. Interannual growth patterns were strongly correlated between top diameter trees and the other classes, and trees of all diameter classes generally responded to climate in the same way, which indicated that it is sufficient to sample only the largest diameter trees in a stand to provide insight into growth–climate relationships. The forecasted growth estimates indicate that future climate warming will negatively impact the productivity of lodgepole pine in the FH, the heart of lodgepole distribution and productivity in Alberta.     

The effects of herbaceous and woody competition on planted white pine in a clearcut site

We investigated the effects of herbaceous and woody vegetation control on the survival and growth of planted eastern white pine (Pinus strobus L.) seedlings through six growing seasons. Herbaceous vegetation control involved the suppression of grasses, forbs, ferns, and low-shrubs, and was maintained for 0, 2, or 4 years after white pine seedlings were planted. Woody control involved the removal of all tall-shrub and deciduous trees, and was conducted at the time of planting, at the end of the second or fifth growing seasons, or not at all. Seedling height and basal diameter responded positively and proportionally to duration of herbaceous vegetation control. Gains associated with woody control were generally not significant unless some degree of herbaceous vegetation control was also conducted. Only herbaceous control increased pine crown closure and rate of crown closure. Herbaceous control and the presence of 5000–15,000 stems per ha of young overtopping aspen were associated with reduced weevil (Pissodes strobi Peck.) injury and increased pine height growth. The study suggests that white pine restoration strategies on clearcut sites should focus on the proactive, early management of understory vegetation and the gradual reduction of overtopping cover from woody vegetation to create a seedling light environment that supports acceptable growth with minimal weevil damage.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.