A mid-century ecological forecast with partitioned uncertainty predicts increases in loblolly pine forest productivity.

Ecological forecasting of forest productivity involves integrating observations into a process-based model and propagating the dominant components of uncertainty to generate probability distributions for future states and fluxes. Here, we develop a forecast for the biomass change in loblolly pine (Pinus taeda) forests of the southeastern United States and evaluate the relative contribution of different forms of uncertainty to the total forecast uncertainty. Specifically, we assimilated observations of carbon and flux stocks and fluxes from sites across the region, including global change experiments, into a forest ecosystem model to calibrate the parameter distributions and estimate the process uncertainty (i.e., model structure uncertainty revealed in the residuals of the calibration). Using this calibration, we forecasted the change in biomass within each 12-digit Hydrologic (H12) unit across the native range of loblolly pine between 2010 and 2055 under the Representative Concentration Pathway 8.5 scenario. Averaged across the region, productivity is predicted to increase by a mean of 31% between 2010 and 2055 with an average forecast 95% quantile interval of ±15 percentage units. The largest increases were predicted in cooler locations, corresponding to the largest projected changes in temperature. The forecasted mean change varied considerably among the H12 units (3-80% productivity increase), but only units in the warmest and driest extents of the loblolly pine range had forecast distributions with probabilities of a decline in productivity that exceeded 5%. By isolating the individual components of the forecast uncertainty, we found that ecosystem model process uncertainty made the largest individual contribution. Ecosystem model parameter and climate model uncertainty had similar contributions to the overall forecast uncertainty, but with differing spatial patterns across the study region. The probabilistic framework developed here could be modified to include additional sources of uncertainty, including changes due to fire, insects, and pests: processes that would result in lower productivity changes than forecasted here. Overall, this study presents an ecological forecast at the ecosystem management scale so that land managers can explicitly account for uncertainty in decision analysis. Furthermore, it highlights that future work should focus on quantifying, propagating, and reducing ecosystem model process uncertainty.

Estimating Stand Height and Tree Density in Pinus taeda plantations using in-situ data, airborne LiDAR and k-Nearest Neighbor Imputation.

Accurate forest inventory is of great economic importance to optimize the entire supply chain management in pulp and paper companies. The aim of this study was to estimate stand dominate and mean heights (HD and HM) and tree density (TD) of Pinus taeda plantations located in South Brazil using in-situ measurements, airborne Light Detection and Ranging (LiDAR) data and the non- k-nearest neighbor (k-NN) imputation. Forest inventory attributes and LiDAR derived metrics were calculated at 53 regular sample plots and we used imputation models to retrieve the forest attributes at plot and landscape-levels. The best LiDAR-derived metrics to predict HD, HM and TD were H99TH, HSD, SKE and HMIN. The Imputation model using the selected metrics was more effective for retrieving height than tree density. The model coefficients of determination (adj.R2) and a root mean squared difference (RMSD) for HD, HM and TD were 0.90, 0.94, 0.38m and 6.99, 5.70, 12.92%, respectively. Our results show that LiDAR and k-NN imputation can be used to predict stand heights with high accuracy in Pinus taeda. However, furthers studies need to be realized to improve the accuracy prediction of TD and to evaluate and compare the cost of acquisition and processing of LiDAR data against the conventional inventory procedures.

Pinus taeda forest growth predictions in the 21st century vary with site mean annual temperature and site quality.

Climate projections from 20 downscaled global climate models (GCMs) were used with the 3-PG model to predict the future productivity and water use of planted loblolly pine (Pinus taeda) growing across the southeastern United States. Predictions were made using Representative Concentration Pathways (RCP) 4.5 and 8.5. These represent scenarios in which total radiative forcing stabilizes before 2100 (RCP 4.5) or continues increasing throughout the century (RCP 8.5). Thirty-six sites evenly distributed across the native range of the species were used in the analysis. These sites represent a range in current mean annual temperature (14.9-21.6°C) and precipitation (1,120-1,680 mm/year). The site index of each site, which is a measure of growth potential, was varied to represent different levels of management. The 3-PG model predicted that aboveground biomass growth and net primary productivity will increase by 10%-40% in many parts of the region in the future. At cooler sites, the relative growth increase was greater than at warmer sites. By running the model with the baseline [CO2 ] or the anticipated elevated [CO2 ], the effect of CO2 on growth was separated from that of other climate factors. The growth increase at warmer sites was due almost entirely to elevated [CO2 ]. The growth increase at cooler sites was due to a combination of elevated [CO2 ] and increased air temperature. Low site index stands had a greater relative increase in growth under the climate change scenarios than those with a high site index. Water use increased in proportion to increases in leaf area and productivity but precipitation was still adequate, based on the downscaled GCM climate projections. We conclude that an increase in productivity can be expected for a large majority of the planted loblolly pine stands in the southeastern United States during this century.

Modeling and mapping basal area of Pinus taeda L. plantation using airborne LiDAR data.

Basal area (BA) is a good predictor of timber stand volume and forest growth. This study developed predictive models using field and airborne LiDAR (Light Detection and Ranging) data for estimation of basal area in Pinus taeda plantation in south Brazil. In the field, BA was collected from conventional forest inventory plots. Multiple linear regression models for predicting BA from LiDAR-derived metrics were developed and evaluated for predictive power and parsimony. The best model to predict BA from a family of six models was selected based on corrected Akaike Information Criterion (AICc) and assessed by the adjusted coefficient of determination (adj. R²) and root mean square error (RMSE). The best model revealed an adj. R²=0.93 and RMSE=7.74%. Leave one out cross-validation of the best regression model was also computed, and revealed an adj. R² and RMSE of 0.92 and 8.31%, respectively. This study showed that LiDAR-derived metrics can be used to predict BA in Pinus taeda plantations in south Brazil with high precision. We conclude that there is good potential to monitor growth in this type of plantations using airborne LiDAR. We hope that the promising results for BA modeling presented herein will stimulate to operate this technology in Brazil.

Climate Change and Ecosystem Services Output Efficiency in Southern Loblolly Pine Forests.

Forests provide myriad ecosystem services that are vital to humanity. With climate change, we expect to see significant changes to forests that will alter the supply of these critical services and affect human well-being. To better understand the impacts of climate change on forest-based ecosystem services, we applied a data envelopment analysis method to assess plot-level efficiency in the provision of ecosystem services in Florida natural loblolly pine (Pinus taeda L.) forests. Using field data for n = 16 loblolly pine forest plots, including inputs such as site index, tree density, age, precipitation, and temperatures for each forest plot, we assessed the relative plot-level production of three ecosystem services: timber, carbon sequestered, and species richness. The results suggested that loblolly pine forests in Florida were largely inefficient in the provision of these ecosystem services under current climatic conditions. Climate change had a small negative impact on the loblolly pine forests efficiency in the provision of ecosystem services. In this context, we discussed the reduction of tree density that may not improve ecosystem services production.

SRWC bioenergy productivity and economic feasibility on marginal lands.

Evolving bioenergy markets necessitate consideration of marginal lands for woody biomass production worldwide particularly the southeastern U.S., a prominent wood pellet exporter to Europe. Growing short rotation woody crops (SRWCs) on marginal lands minimizes concerns about using croplands for bioenergy production and reinforces sustainability of wood supply to existing and growing global biomass markets. We estimated mean annual aboveground green biomass increments (MAIs) and assessed economic feasibility of various operationally established (0.5 ha-109 ha) SRWC stands on lands used to mitigate environmental liabilities of municipal wastewater, livestock wastewater and sludge, and subsurface contamination by petroleum and pesticides. MAIs (Mg ha(-1) yr(-1)) had no consistent relationship with stand density or age. Non-irrigated Populus, Plantanus occidentalis L. and Pinus taeda L. stands produced 2.4-12.4 Mg ha(-1) yr(-1). Older, irrigated Taxodium distchum L., Fraxinus pennsylvanica L., and coppiced P. occidentalis stands had higher MAIs (10.6-21.3 Mg ha(-1) yr(-1)) than irrigated Liquidambar styraciflua L. and non-coppiced, irrigated P. occidentalis (8-18 Mg ha(-1) yr(-1)). Natural hardwood MAIs at 20-60 years were less than hardwood and P. taeda productivities at 5-20 years. Unlike weed control, irrigation and coppicing improved managed hardwood productivity. Rotation length affected economic outcomes although the returns were poor due to high establishment and maintenance costs, low productivities and low current stumpage values, which are expected to quickly change with development of robust global markets.

Understanding the dynamic evolution of hemicellulose during Pinus taeda L. growth.

Pinus taeda L. is a fast-growing softwood with significant commercial value. Understanding structural changes in hemicellulose during growth is essential to understanding the biosynthesis processes occurring in the cell walls of this tree. In this study, alkaline extraction is applied to isolate hemicellulose from Pinus taeda L. stem segments of different ages (1, 2, 3, and 4 years old). The results show that the extracted hemicellulose is mainly comprised of O-acetylgalactoglucomannan (GGM) and 4-O-methylglucuronoarabinoxylan (GAX), with the molecular weights and ratios (i.e., GGM:GAX) of GGM and GAX increasing alongside Pinus taeda L. age. Mature Pinus taeda L. hemicellulose is mainly composed of GGM, and the ratio of (mannose:glucose) in the GGM main chain gradually increases from 2.45 to 3.60 with growth, while the galactose substitution of GGM decreases gradually from 21.36% to 14.65%. The acetylation of GGM gradually increases from 0.33 to 0.45 with the acetyl groups mainly substituting into the O-3 position in the mannan. Furthermore, the contents of arabinose and glucuronic acid in GAX gradually decrease with growth. This study can provide useful information to the research in genetic breeding and high-value utilization of Pinus taeda L.

Association genetics of oleoresin flow in loblolly pine: discovering genes and predicting phenotype for improved resistance to bark beetles and bioenergy potential.

Rapidly enhancing oleoresin production in conifer stems through genomic selection and genetic engineering may increase resistance to bark beetles and terpenoid yield for liquid biofuels. We integrated association genetic and genomic prediction analyses of oleoresin flow (g 24 h(-1)) using 4854 single nucleotide polymorphisms (SNPs) in expressed genes within a pedigreed population of loblolly pine (Pinus taeda) that was clonally replicated at three sites in the southeastern United States. Additive genetic variation in oleoresin flow (h(2) ≈ 0.12-0.30) was strongly correlated between years in which precipitation varied (r(a) ≈ 0.95), while the genetic correlation between sites declined from 0.8 to 0.37 with increasing differences in soil and climate among sites. A total of 231 SNPs were significantly associated with oleoresin flow, of which 81% were specific to individual sites. SNPs in sequences similar to ethylene signaling proteins, ABC transporters, and diterpenoid hydroxylases were associated with oleoresin flow across sites. Despite this complex genetic architecture, we developed a genomic prediction model to accelerate breeding for enhanced oleoresin flow that is robust to environmental variation. Results imply that breeding could increase oleoresin flow 1.5- to 2.4-fold in one generation.

On the complementary relationship between marginal nitrogen and water-use efficiencies among Pinus taeda leaves grown under ambient and CO2-enriched environments.

BACKGROUND AND AIMS: Water and nitrogen (N) are two limiting resources for biomass production of terrestrial vegetation. Water losses in transpiration (E) can be decreased by reducing leaf stomatal conductance (g(s)) at the expense of lowering CO(2) uptake (A), resulting in increased water-use efficiency. However, with more N available, higher allocation of N to photosynthetic proteins improves A so that N-use efficiency is reduced when g(s) declines. Hence, a trade-off is expected between these two resource-use efficiencies. In this study it is hypothesized that when foliar concentration (N) varies on time scales much longer than g(s), an explicit complementary relationship between the marginal water- and N-use efficiency emerges. Furthermore, a shift in this relationship is anticipated with increasing atmospheric CO(2) concentration (c(a)). METHODS: Optimization theory is employed to quantify interactions between resource-use efficiencies under elevated c(a) and soil N amendments. The analyses are based on marginal water- and N-use efficiencies, λ = (∂A/∂g(s))/(∂E/∂g(s)) and η = ∂A/∂N, respectively. The relationship between the two efficiencies and related variation in intercellular CO(2) concentration (c(i)) were examined using A/c(i) curves and foliar N measured on Pinus taeda needles collected at various canopy locations at the Duke Forest Free Air CO(2) Enrichment experiment (North Carolina, USA). KEY RESULTS: Optimality theory allowed the definition of a novel, explicit relationship between two intrinsic leaf-scale properties where η is complementary to the square-root of λ. The data support the model predictions that elevated c(a) increased η and λ, and at given c(a) and needle age-class, the two quantities varied among needles in an approximately complementary manner. CONCLUSIONS: The derived analytical expressions can be employed in scaling-up carbon, water and N fluxes from leaf to ecosystem, but also to derive transpiration estimates from those of η, and assist in predicting how increasing c(a) influences ecosystem water use.

Soil fungal cellobiohydrolase I gene (cbhI) composition and expression in a loblolly pine plantation under conditions of elevated atmospheric CO2 and nitrogen fertilization.

The simultaneous increase of atmospheric CO(2) and nitrogen (N) deposition to terrestrial ecosystems is predicted to alter plant productivity and, consequently, to change the amount and quality of above- and belowground carbon entering forest soils. It is not known how such changes will impact the composition and function of soil fungal communities that play a key role in degrading complex carbon. We sequenced the fungal cellobiohydrolase I gene (cbhI) from soil DNA and cDNA to compare the richness and composition of resident and expressed cbhI genes at a U.S. Department of Energy free air-carbon dioxide enrichment (FACE) site (NC), which had been exposed to elevated atmospheric CO(2) and/or N fertilization treatment for several years. Our results provide evidence that the richness and composition of the cellulolytic fungi surveyed in this study were distinct in the DNA- and cDNA-based gene surveys and were dominated by Basidiomycota that have low or no representation in public databases. The surveys did not detect differences in richness or phylum-level composition of cbhI-containing, cellulolytic fungi that correlated with elevated CO(2) or N fertilization at the time of sampling.

Myo-inositol hexakisphosphate, isolated from female gametophyte tissue of loblolly pine, inhibits growth of early-stage somatic embryos.

• Myo-inositol hexakisphosphate (InsP(6)), abundant in animals and plants, is well known for its anticancer activity. However, many aspects of InsP(6) function in plants remain undefined. We now report the first evidence that InsP(6) can inhibit cellular proliferation in plants under growth conditions where phosphorus is not limited. • A highly anionic molecule inhibitory to early-stage somatic embryo growth of loblolly pine (LP) was purified chromatographically from late-stage LP female gametophytes (FGs), and then characterized structurally using mass spectrometry (MS) and nuclear magnetic resonance (NMR) analyses. • Exact mass and mass spectrometry-mass spectrometry (MS-MS) fragmentation identified the bioactive molecule as an inositol hexakisphosphate. It was then identified as the myo-isomer (i.e. InsP(6)) on the basis of (1)H-, (31)P- and (13)C-NMR, (1)H-(1)H correlation spectroscopy (COSY), (1)H-(31)P heteronuclear single quantum correlation (HSQC) and (1)H-(13)C HSQC. Topical application of InsP(6) to early-stage somatic embryos indeed inhibits embryonic growth. • Recently evidence has begun to emerge that InsP(6) may also play a regulatory role in plant cells. We anticipate that our findings will help to stimulate additional investigations aimed at elucidating the roles of inositol phosphates in cellular growth and development in plants.

Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation.

The degree to which rising atmospheric CO(2) will be offset by carbon (C) sequestration in forests depends in part on the capacity of trees and soil microbes to make physiological adjustments that can alleviate resource limitation. Here, we show for the first time that mature trees exposed to CO(2) enrichment increase the release of soluble C from roots to soil, and that such increases are coupled to the accelerated turnover of nitrogen (N) pools in the rhizosphere. Over the course of 3 years, we measured in situ rates of root exudation from 420 intact loblolly pine (Pinus taeda L.) roots. Trees fumigated with elevated CO(2) (200 p.p.m.v. over background) increased exudation rates (µg C cm(-1) root h(-1) ) by 55% during the primary growing season, leading to a 50% annual increase in dissolved organic inputs to fumigated forest soils. These increases in root-derived C were positively correlated with microbial release of extracellular enzymes involved in breakdown of organic N (R(2) = 0.66; P = 0.006) in the rhizosphere, indicating that exudation stimulated microbial activity and accelerated the rate of soil organic matter (SOM) turnover. In support of this conclusion, trees exposed to both elevated CO(2) and N fertilization did not increase exudation rates and had reduced enzyme activities in the rhizosphere. Collectively, our results provide field-based empirical support suggesting that sustained growth responses of forests to elevated CO(2) in low fertility soils are maintained by enhanced rates of microbial activity and N cycling fuelled by inputs of root-derived C. To the extent that increases in exudation also stimulate SOM decomposition, such changes may prevent soil C accumulation in forest ecosystems.

Relationships between stem CO(2) efflux, substrate supply, and growth in young loblolly pine trees.

*We examined the relationships between stem CO(2) efflux (E(s)), diameter growth, and nonstructural carbohydrate concentration in loblolly pine trees. Carbohydrate supply was altered via stem girdling during rapid stem growth in the spring and after growth had ceased in the autumn. We hypothesized that substrate type and availability control the seasonal variation and temperature sensitivity of E(s). *The E(s) increased and decreased above and below the girdle, respectively, within 24 h of treatment. Seasonal variation in E(s) response to girdling corresponded to changes in stem soluble sugar and starch concentration. Relative to nongirdled trees, E(s) increased 94% above the girdle and decreased 50% below in the autumn compared with a 60% and 20% response at similar positions in the spring. *The sensitivity of E(s) to temperature decreased below the girdle in the autumn and spring and increased above the girdle but only in the autumn. Temperature-corrected E(s) was linearly related to soluble sugar (R(2) = 0.57) and starch (R(2) = 0.62) concentration. *We conclude that carbohydrate supply, primarily recently fixed photosynthate, strongly influences E(s) in Pinus taeda stems. Carbohydrate availability effects on E(s) obviate the utility of applying short-term temperature response functions across seasons.

Soil incorporation of logging residue affects fine-root and mycorrhizal root-tip dynamics of young loblolly pine clones.

Loblolly pine (Pinus taeda L.) plantations cover a large geographic area of the southeastern USA and supply a large proportion of the nation's wood products. Research on management strategies designed to maximize wood production while also optimizing nutrient use efficiency and soil C sequestration is needed. We used minirhizotrons to quantify the effects of incorporating logging residues into soil on fine-root standing crop, production and mortality, and mycorrhizal root tips in young loblolly pine clones of contrasting ideotypes. Clone 93 is known to allocate more C to stem growth, while clone 32 allocates less C to stems and more to leaves. The relative allocation by these clones to support fine-root turnover is unknown. Clone 32 exhibited 37% more fine-root mortality than clone 93, which was mainly the result of a greater standing crop of fine roots. Fine-root standing crop in plots amended with logging residue was initially higher than control plots, but 2.5 years after planting, standing crop in control plots had exceeded that in mulched plots. Production of mycorrhizal root tips, on the other hand, was initially higher in control than mulched plots, but during the last 9 months of the study, mycorrhizal tip production was greater in mulched than control plots, especially for clone 93. As expected, turnover rate of fine roots was greater in surface soil (0-25 cm) compared with deeper (25-50 cm) soil and for small roots (< 0.4 mm diameter) compared with larger fine roots (0.4-2.0 mm diameter). Rates of fine-root turnover were similar in both clones. Organic matter additions reduced survivorship of individual roots and increased turnover rates of fine-root populations. Results indicate that management decisions should be tailored to fit the growth and allocation patterns of available clones.

Identification and characterization of a matrix metalloproteinase (Pta1-MMP) expressed during Loblolly pine (Pinus taeda) seed development, germination completion, and early seedling establishment.

Extracellular matrix (ECM) modifications occur during plant growth, development, and in response to environmental stimuli. Key modulators of ECM modification in vertebrates, the extracellular matrix metalloproteinases (MMPs), have also been described in a few plants. Here, we report the identification of Loblolly pine (Pinus taeda) Pta1-MMP and its characterization during seed development and germination. Pta1-MMP protein has the structural characteristics of other plant MMPs, the recombinant protein exhibits Zn(2+)-dependent protease activity, and is inhibited by EDTA and the active site-binding hydroxamate inhibitor GM6001. The Pta1-MMP gene is expressed in both embryo and megagametophyte, with transcript levels increasing in both during the period from proembryo to early cotyledonary stage, then declining during late embryogenesis and maturation drying. Protein extracts exhibited similar developmental-stage MMP-like activity. Seed germination was stimulated by GA(3) and inhibited by ABA, and the timing of germination completion was mirrored by the presence of MMP-like protease activity in both water- and GA(3)-imbibed embryos. Pta1-MMP gene transcript levels increased in association with radicle protrusion for both GA(3)- and water-treated embryos, in agreement with MMP-like activity. In contrast, by 11 days after imbibition, Pta1-MMP gene transcripts in ABA-treated embryos were at levels similar to the other treatments, although MMP-like activity was not observed. The application of GM6001 during Loblolly pine seed germination inhibited radicle protrusion. Our results suggest that MMP activity may be involved in ECM modification, facilitating the cell division and expansion required during seed development, germination completion, and subsequent seedling establishment.

Predicting the regenerative capacity of conifer somatic embryogenic cultures by metabolomics.

Somatic embryogenesis in gymnosperms is an effective approach to clonally propagating germplasm. However, embryogenic cultures frequently lose regenerative capacity. The interactions between metabolic composition, physiological state, genotype and embryogenic capacity in Pinus taeda (loblolly pine) somatic embryogenic cultures were explored using metabolomics. A stepwise modelling procedure, using the Bayesian information criterion, generated a 47 metabolite predictive model that could explain culture productivity. The model performed extremely well in cross-validation, achieving a correlation coefficient of 0.98 between actual and predicted mature embryo production. The metabolic composition and structure of the model implied that variation in culture regenerative capacity was closely linked to the physiological transition of cultures from the proliferation phase to the maturation phase of development. The propensity of cultures to advance into this transition appears to relate to nutrient uptake and allocation in vivo, and to be associated with the tolerance and response of cultures to stress, during the proliferation phase.

Acclimation of leaf hydraulic conductance and stomatal conductance of Pinus taeda (loblolly pine) to long-term growth in elevated CO(2) (free-air CO(2) enrichment) and N-fertilization.

We investigated how leaf hydraulic conductance (K(leaf)) of loblolly pine trees is influenced by soil nitrogen amendment (N) in stands subjected to ambient or elevated CO(2) concentrations (CO(2)(a) and CO(2)(e), respectively). We also examined how K(leaf) varies with changes in reference leaf water potential (Psi(leaf-ref)) and stomatal conductance (g(s-ref)) calculated at vapour pressure deficit, D of 1 kPa. We detected significant reductions in K(leaf) caused by N and CO(2)(e), but neither treatment affected pre-dawn or midday Psi(leaf). We also detected a significant CO(2)(e)-induced reduction in g(s-ref) and Psi(leaf-ref). Among treatments, the sensitivity of K(leaf) to Psi(leaf) was directly related to a reference K(leaf) (K(leaf-ref) computed at Psi(leaf-ref)). This liquid-phase response was reflected in a similar gas-phase response, with g(s) sensitivity to D proportional to g(s-ref). Because leaves represented a substantial component of the whole-tree conductance, reduction in K(leaf) under CO(2)(e) affected whole-tree water use by inducing a decline in g(s-ref). The consequences of the acclimation of leaves to the treatments were: (1) trees growing under CO(2)(e) controlled morning leaf water status less than CO(2)(a) trees resulting in a higher diurnal loss of K(leaf); (2) the effect of CO(2)(e) on g(s-ref) was manifested only during times of high soil moisture.

Loblolly pine (Pinus taeda) female gametophyte and embryo pH changes during seed development.

Stage-specific measurements of female gametophyte (FG) and embryo pH (hydrogen ion concentration) were made through the sequence of loblolly pine (Pinus taeda L.) seed development. The FG tissue from two open-pollinated trees showed similar pH profiles starting at 5.5 shortly after fertilization, increasing to about 6.1 at stage 7, levelling off at 6.3-6.5 towards the end of development and dropping to 6.0 just before cone opening. Measurements of the chalazal end were 0.05-0.2 pH units less than the micropylar end through early-to-mid-development. In contrast, embryo pH maintained a nearly constant value near 7.0 through development. Profiles of pH through seed development were similar whether portrayed by date or stage of embryo present in the seed. The pH profiles assisted in the development of improved embryogenic tissue initiation techniques. When post-autoclaving maturation medium pH was raised from about 5.3 in control medium to 5.7 or 5.5-5.7 with 2(n-morpholino)ethanesulphonic acid, cotyledonary embryo yields increased.

Osmotic measurements in whole megagametophytes and embryos of loblolly pine (Pinus taeda) during seed development.

Water potential (Psi) and osmotic potential (Psis) were measured weekly through the sequence of seed development in megagametophytes of loblolly pine (Pinus taeda L.). A Wescor 5500XRS vapor pressure osmometer, modified with a cycle hold switch, was used to measure Psi for whole megagametophytes containing embryos. The Psi measurements for megagametophytes with embryos removed were also attempted but readings were distorted due to cell lysates from the cut surfaces. Six seasonal sets of megagametophyte Psi profiles were generated. Megagametophytes from most of the trees examined showed a consistent Psi pattern: low measurements of -1.0 to -0.75 MPa during early embryo development in late June to early July when embryo Stages 1-2 occur; an increase for one to several weeks to levels of -0.5 to -0.75 MPa, beginning at Stages 3-5 when apical dome formation occurs; followed by a steady drop from -0.85 to -1.7 to -2.0 MPa from Stage 6 onward from late August until just before cone seed release. The Psis was measured for supernatant from centrifuged frozen-thawed megagametophyte tissue (embryos removed). Megagametophyte Psis profiles were similar for seeds analyzed from two trees and resembled Psi observations starting low, rising around Stages 4-7 and then undergoing a major reduction indicating a strong solute accumulation beginning at Stages 7-9.1. Somatic embryos stop growth prematurely in vitro at Stages 8-9.1. The major change in the accumulation of megagametophyte solutes at Stages 8-9.1 correlates with the halt in somatic embryo maturation and suggests that identifying, quantifying and using the major natural soluble compounds that accumulate during mid- to late-stage seed development may be important to improve conifer somatic embryo maturation.

Pinus taeda clones and soil nutrient availability: effects of soil organic matter incorporation and fertilization on biomass partitioning and leaf physiology.

The combined effects of intensive management and planting of improved seedlings have led to large increases in productivity on intensively managed pine forests in the southeastern United States. To best match clones to particular site conditions, an understanding of how specific clones respond to changes in nutrition in terms of biomass partitioning, leaf physiology and biochemistry will be necessary. This study measured the response of biomass partitioning, light-saturated net photosynthesis (A(Sat)) and photosynthetic capacity to a range in soil fertility and fertilization between two contrasting Pinus taeda L. clone ideotypes: a 'narrow crown' clone (NC) that allocates more resources to stem growth and a 'broad crown' clone (BC) that allocates more resources to leaf area (LA). Under field conditions, we found consistent clone by environment (i.e., varying nutrient regimes) interactions in biomass as well as leaf physiology. Nutrient limitations induced by logging residue incorporation resulted in a 25% loss in stem growth in BC, while NC showed no response. We postulated that the decrease in BC was due to the differences in canopy architecture leading to a reduced canopy CO(2) assimilation, as well as to increased belowground maintenance costs associated with fine-root production. In contrast, N and P additions resulted in a 21% greater increase in stem volume in NC relative to BC. Fertilization increased A(Sat) temporarily in both clones, but A(Sat) eventually decreased below control levels by the end of the study. Although we found a clone by fertilization interaction in leaf physiology, the greatest genotype by environment interaction was found in the LA that appeared to have a greater influence than A(Sat) on growth. This research demonstrates the potential importance of selecting appropriate clonal material and silvicultural prescription when implementing site-specific silviculture to maximize productivity in intensively managed southern pine forests.