Summary• Availability of growth limiting resources may alter root dynamics in forest ecosystems, possibly affecting the land-atmosphere exchange of carbon. This was evaluated for a commercially important southern timber species by installing a factorial experiment of fertilization and irrigation treatments in an 8-yr-old loblolly pine ( Pinus taeda ) plantation.• After 3 yr of growth, production and turnover of fine, coarse and mycorrhizal root length was observed using minirhizotrons, and compared with stem growth and foliage development.• Fertilization increased net production of fine roots and mycorrhizal roots, but did not affect coarse roots. Fine roots had average lifespans of 166 d, coarse roots 294 d and mycorrhizal roots 507 d. Foliage growth rate peaked in late spring and declined over the remainder of the growing season, whereas fine roots experienced multiple growth flushes in the spring, summer and fall.• We conclude that increased nutrient availability might increase carbon input to soils through enhanced fine root turnover. However, this will depend on the extent to which mycorrhizal root formation is affected, as these mycorrhizal roots have much longer average lifespans than fine and coarse roots.
The growth of many pine plantations in the southern United States is limited by soil nutrient availability. Therefore, forest fertilization is a common silvicultural practice throughout the South. Approximately 1.2 million ac of pine plantations were fertilized in 2004. In the last 10 years, considerable advances have been made in identifying the ecophysiological basis for stand growth and the response to fertilizer additions. Nitrogen (N) and phosphorus (P) are the nutrients that most commonly limit growth of southern pine. On wet clay soils in the lower Coastal Plain and on some well-drained soil in the upper Coastal Plain, severe P deficiencies exist. On these soils, P fertilization with 25–50 lb of P per acre at the time of planting produces a large and sustained growth response, on the order of 50 ft3 ac−1 yr−1 (1.5 tn ac−1 yr−1) throughout the rotation. On most other soils in the South, chronic deficiencies of both N and P exist. On these sites, soil nutrient availability often is adequate early in the rotation when tree demand is small. However, around the time of crown closure, N and P frequently become limiting. Fertilization with both N and P in these intermediate aged stands typically increases growth for 8–10 years. The growth response to a combination of 25 lb of P per acre plus 200 lb of N per acre averages around 55 ft3 ac−1 yr−1 (1.6 tn ac−1 yr−1) for an 8-year period. The amount of leaf area in the stand is the main factor determining the current growth rate of the stand and the potential growth response after fertilization. When stand leaf area index is less than 3.5, light capture by the stand is restricted and growth is negatively affected. In many of these stands, fertilization will increase leaf area because of increased soil nutrient availability and thus increase growth. The financial return after fertilization depends on the growth response that occurs, the cost of the fertilizer treatment, and the stumpage value of the timber produced. Using a growth response of 55 ft3 ac−1 yr−1 over 8 years, a fertilizer cost of $90 ac−1, and stumpage values from the first quarter of 2006, the internal rate of return from midrotation fertilization of a loblolly pine plantation with N and P would be approximately 16%.
Changing environmental conditions have the potential to alter allometric relationships between plant parts, possibly leading to ecosystem-level feedbacks. We quantified allometric shifts in field-grown loblolly pine (Pinus taeda L.) in response to altered resource availability based on data from multiple harvests to correct for size-related changes in biomass partitioning. A replicated factorial arrangement of irrigation and fertilization treatments was applied for 4 years to an 8-year-old loblolly pine plantation on a well-drained, low fertility site in North Carolina. Destructive and nondestructive growth measurements were used to develop treatment-specific regressions to estimate stand-level biomass for ephemeral and perennial plant parts, both above- and belowground. Stand-level allometric analysis indicated that irrigation increased biomass partitioning to fine roots and decreased partitioning to foliage, relative to other plant parts. Fertilization increased partitioning to perennial tissues (coarse roots, taproots, and branches) and decreased partitioning to ephemeral tissues (foliage and fine roots). Changes in allometry were small (< 6 %) but statistically significant, indicating that biomass partitioning in loblolly pine changes with altered resource availability, but is probably under strong ontogenetic control.
We used estimates of autotrophic respiration (R A ), net primary productivity (NPP) and soil CO 2 evolution (S ff ), to develop component carbon budgets for 12-year-old loblolly pine plantations during the fifth year of a fertilization and irrigation experiment. Annual carbon use in R A was 7.5, 9.0, 15.0, and 15.1 Mg C ha À1 in control (C), irrigated (I), fertilized (F) and irrigated and fertilized (IF) treatments, respectively. Foliage, fine root and perennial woody tissue (stem, branch, coarse and taproot) respiration accounted for, respectively, 37%, 24%, and 39% of R A in C and I treatments and 38%, 12% and 50% of R A in F and IF treatments. Annual gross primary production (GPP 5 NPP 1 R A ) ranged from 13.1 to 26.6 Mg C ha À1 . The I, F, and IF treatments resulted in a 21, 94, and 103% increase in GPP, respectively, compared to the C treatment. Despite large treatment differences in NPP, R A , and carbon allocation, carbon use efficiency (CUE 5 NPP/GPP) averaged 0.42 and was unaffected by manipulating site resources.Ecosystem respiration (R E ), the sum of S ff , and above ground R A , ranged from 12.8 to 20.2 Mg C ha À1 yr À1 . S ff contributed the largest proportion of R E , but the relative importance of S ff decreased from 0.63 in C treatments to 0.47 in IF treatments because of increased aboveground R A . Aboveground woody tissue R A was 15% of R E in C and I treatments compared to 25% of R E in F and IF treatments. Net ecosystem productivity (NEP 5 GPP-R E ) was roughly 0 in the C and I treatments and 6.4 Mg C ha À1 yr À1 in F and IF treatments, indicating that non-fertilized treatments were neither a source nor a sink for atmospheric carbon while fertilized treatments were carbon sinks. In these young stands, NEP is tightly linked to NPP; increased ecosystem carbon storage results mainly from an increase in foliage and perennial woody biomass.
Abstract. Predicting how forest carbon cycling will change in response to climate change and management depends on the collective knowledge from measurements across environmental gradients, ecosystem manipulations of global change factors, and mathematical models. Formally integrating these sources of knowledge through data assimilation, or model-data fusion, allows the use of past observations to constrain model parameters and estimate prediction uncertainty. Data assimilation (DA) focused on the regional scale has the opportunity to integrate data from both environmental gradients and experimental studies to constrain model parameters. Here, we introduce a hierarchical Bayesian DA approach (Data Assimilation to Predict Productivity for Ecosystems and Regions, DAPPER) that uses observations of carbon stocks, carbon fluxes, water fluxes, and vegetation dynamics from loblolly pine plantation ecosystems across the southeastern US to constrain parameters in a modified version of the Physiological Principles Predicting Growth (3-PG) forest growth model. The observations included major experiments that manipulated atmospheric carbon dioxide (CO 2 ) concentration, water, and nutrients, along with nonexperimental surveys that spanned environmental gradients across an 8.6 × 10 5 km 2 region. We optimized regionally representative posterior distributions for model parameters, which dependably predicted data from plots withheld from the data assimilation. While the mean bias in predictions of nutrient fertilization experiments, irrigation experiments, and CO 2 enrichment experiments was low, future work needs to focus modifications to model structures that decrease the bias in predictions of drought experiments. Predictions of how growth responded to elevated CO 2 strongly depended on whether ecosystem experiments were assimilated and whether the assimilated field plots in the CO 2 study were allowed to have different mortality parameters than the other field plots in the region. We present predictions of stem biomass productivity under elevated CO 2 , decreased precipPublished by Copernicus Publications on behalf of the European Geosciences Union. itation, and increased nutrient availability that include estimates of uncertainty for the southeastern US. Overall, we (1) demonstrated how three decades of research in southeastern US planted pine forests can be used to develop DA techniques that use multiple locations, multiple data streams, and multiple ecosystem experiment types to optimize parameters and (2) developed a tool for the development of future predictions of forest productivity for natural resource managers that leverage a rich dataset of integrated ecosystem observations across a region.
A study of the effects of nutrients and water supply (2 ϫ 2 factorial experiment) was conducted in a 12-yr-old stand of loblolly pine (Pinus taeda L.) during a period in which soil moisture was not augmented by irrigation because of frequent rain events. Information on the responses of sapwood-to-leaf area ratio and early-to-late wood ratio, to four years of treatments led to the hypothesis that the combination of increased nutrient and water supply (IF treatment) will increase tree transpiration rate per unit leaf area (E C,1 ) above E C,1 in the control (C), as well as increasing E C,1 above that when either the supply of water (I) or of nutrients (F) is increased. We further hypothesized that canopy transpiration (E C ) will rank IF Ͼ F Ͼ I ϭ C, based on the ranking of leaf area index (L) and assuming that the ranking of E C,1 is as first hypothesized. We rejected our first hypothesis, because F had lower E C,1 than the other treatments, rather than IF having higher values. We could not reject the second hypothesis; the ranking of average daily E C was 1.8 mm for IF, 1.2 mm for F, and 0.7 mm for both C and I (SE Ͻ 0.1 mm for all treatments). Thus, it was the lower E C,1 of the F treatment, relative to IF, that resulted in ranking of E C similar to that hypothesized. Lower E C,1 in F trees was found to relate to lower canopy stomatal conductance, even though soil moisture conditions during the time of the study were similar in all treatments. Only trees in the F treatment absorbed a substantial amount of water (25%) below 1 m in the soil. These results indicate a ''carry-over'' effect of irrigation when combined with fertilization that increases E C in irrigated trees, relative to unirrigated trees, even under conditions when soil moisture is high and similar in all treatments.
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