Intra-annual nutrient flux in Pinus taeda.

Intra-annual nutrient (nitrogen, phosphorus, potassium, calcium and magnesium) flux was quantified for Pinus taeda L. at a nutrient-poor, well-drained sandy site in Scotland County, NC, USA where a 2 × 2 factorial of irrigation and nutrition was applied in four replications in a 10-year-old stand with 1200 stems ha(-1). Treatments were applied with the goal of providing optimum nutrition (no nutritional deficiencies) and water availability. Component (foliage, branch, stem and root) nutrient content was estimated monthly for 2 years using nutrient concentration and phenology assessments combined with destructive harvests. Positive flux values indicated nutrient accumulation in the trees while negative values indicated nutrient loss from the trees. Fertilization significantly increased nitrogen, phosphorus, potassium, calcium and magnesium flux 140%, on average, over non-fertilized. Irrigation significantly increased calcium flux 28% while there was no significant irrigation effect on nitrogen, phosphorus, potassium or magnesium. Maximum nutrient fluxes (kg ha(-1) day(-1)) for non-fertilized and fertilized stands were 0.36 and 1.05 for nitrogen, 0.042 and 0.095 for phosphorus, 0.13 and 0.51 for potassium, 0.27 and 0.42 for calcium, and 0.04 and 0.12 for magnesium, respectively. Maximum flux was coincident with ephemeral tissue (foliage and fine root) development and likely would be higher in stands with more foliage than those observed in this study (projected leaf area indices were 1.5 and 3.0 for the non-fertilized and fertilized stands). Minimum nutrient fluxes (kg ha(-1) day(-1)) for non-fertilized and fertilized stands were -0.18 and -0.42 for nitrogen, -0.029 and -0.070 for phosphorus, -0.05 and -0.18 for potassium, -0.04 and -0.05 for calcium, and -0.02 and -0.03 for magnesium, respectively. Minimum fluxes were typically observed in the dormant season and were linked to foliage senescence and branch death. Foliage and branch component nutrient contents were out of phase for nitrogen, phosphorus, potassium and magnesium, indicating nutrient retranslocation and storage in branches prior to foliage development and after foliage senescence. In contrast to current operational fertilizer programs which often target winter application these data suggest the best application times would be during foliage development.

Nutrient use and uptake in Pinus taeda.

We quantified nitrogen (N), phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) content, use (nutrient amount for one growth year), retranslocation (nutrients recycled before foliage senescence), uptake (use minus retranslocation), volume production per unit of uptake and fertilizer-uptake efficiency (percent applied taken up) in a 2 x 2 (nutrient and water) factorial experiment replicated four times in an 8-year-old loblolly pine (Pinus taeda L.) stand growing on a nutrient-poor sandy soil in Scotland County, North Carolina, USA. Over 14 years, we applied 1140, 168, 393, 168 and 146 kg ha(-1) of elemental N, P, K, Ca and Mg fertilizer, respectively, and an average of 710 mm year(-1) of irrigation. All plots received complete vegetation control. Fertilization about doubled tissue N, P, K and Mg contents at age 21, whereas irrigation resulted in smaller increases in nutrient contents. Maximum annual uptake was 101, 9.3, 44, 37 and 13 kg ha(-1) year(-1) and volume production per unit of nutrient uptake was 0.35, 3.5, 0.66, 1.1 and 3.1 m(3) kg(-1), for N, P, K, Ca and Mg, respectively. Irrigated plots had greater volume production per unit of N, P, K and Mg uptake than control plots, likely because irrigation allowed photosynthesis to continue during dry periods. Fertilized plus irrigated plots had less volume production per unit of these elements than the fertilized plots either because nutrient uptake exceeded the requirement for optimum growth or because available water (rainfall plus irrigation) was insufficient for the leaf area achieved with fertilization. At age 19, fertilizer-uptake efficiencies for N, P, K, Ca and Mg were 53, 24, 62, 57 and 39%, respectively, and increased with irrigation to 68, 36, 78, 116 and 55%, respectively. The scale of fertilizer uptake was likely a result of low native site nutrient availability, study longevity, measurement of all tissue components on site, a comprehensive assessment of coarse roots, and the 3-m rooting depth. Ecosystem nitrogen retention (applied nitrogen found in living plant material, litter fall and soil to 150-cm depth) was estimated at 79% at age 17, a value that would likely be greater when including soil nitrogen to rooting depth and calculating retention at age 21 when the study ended. The ecosystem retention value provides evidence that intensive site resource management can be accomplished with low likelihood of applied materials moving offsite.

Bioavailability of slowly cycling soil phosphorus: major restructuring of soil P fractions over four decades in an aggrading forest.

Although low solubility and slow cycling control P circulation in a wide range of ecosystems, most studies that evaluate bioavailability of soil P use only indices of short-term supply. The objective here is to quantify changes in P fractions in an Ultisol during the growth of an old-field pine forest from 1957 to 2005, specifically changes with organic P (Po) and with inorganic P (Pi) associated with Fe and Al oxides as well as Ca compounds. Changes in soil P were estimated from archived mineral soil samples collected in 1962 shortly after pine seedlings were planted, and on six subsequent occasions (1968, 1977, 1982, 1990, 1997, and 2005) from eight permanent plots and four mineral soil layers (0-7.5, 7.5-15, 15-35, and 35-60 cm). Despite the net transfer of 82.5 kg ha(-1) of P from mineral soil into tree biomass and O horizons, labile soil P was not diminished, as indexed by anion exchange resins, and NaHCO(3) and Mehlich III extractants. An absence of depletion in most labile P fractions masks major restructuring of soil P chemistry driven by ecosystem development. During 28 years of forest growth, decreases were significant and substantial in slowly cycling Po and Pi associated with Fe and Al oxides and Ca compounds, and these accounted for most of the P supplied to biomass and O horizons, and for buffering labile soil fractions as well. Changes in soil P are attributed to the P sink strength of the aggrading forest (at 2.9 kg ha(-1) year(-1) over 28 years); legacies of fertilization, which enriched slowly cycling fractions of Po and Pi; and the changing biogeochemistry of the soil itself.

Effects of mass inoculation on induced oleoresin response in intensively managed loblolly pine.

Oleoresin flow is an important factor in the resistance of pines to attack by southern pine beetle, Dendroctonus frontalis Zimm., and its associated fungi. Abiotic factors, such as nutrient supply and water relations, have the potential to modify this plant-insect-fungus interaction; however, little is known of the effects of inoculation with beetle-associated fungi on oleoresin flow. We observed that constitutive and induced resin yield in loblolly pine, Pinus taeda L., were affected by either fungal inoculation (with the southern pine beetle-associated fungus Ophiostoma minus (Hedgcock) H. & P. Sydow) or silvicultural treatment. The effects of mass wounding (400 wounds m(-2)) and mass wounding and inoculation with O. minus were assessed by comparison with untreated (control) trees. The treatments were applied to trees in a 2 x 2 factorial combination of fertilizer and irrigation treatments. Fertilization did not significantly affect constitutive resin yield. Even as long as 105 days post-treatment, however, mass-inoculated trees produced higher induced resin yields than control or wounded-only trees, indicating a localized induced response to fungal inoculation. We noted no systemic induction of host defenses against fungal colonization. Although beetles attacking previously attacked trees face a greater resinous response from their host than beetles attacking trees that had not been previously attacked, the effect of an earlier attack may not last more than one flight season. Despite mass inoculations, O. minus did not kill the host trees, suggesting that this fungus is not a virulent plant pathogen.

Below-ground carbon input to soil is controlled by nutrient availability and fine root dynamics in loblolly pine.

• 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.

Mineral nutrition, resin flow and phloem phytochemistry in loblolly pine.

Southern pine beetles and associated pathogenic fungi represent the largest biotic threat to pine forests in the southeastern USA. The two primary defensive mechanisms of the tree to the beetle-fungal complex are the primary oleoresin flow and the concentrations of preformed and induced secondary compounds. We compared oleoresin flow and concentrations of phloem nutrients, soluble sugars, starch, total phenolics and proanthocyanidins in Pinus taeda L. trees in fertilized and control plots in the Sandhills region of North Carolina. Four blocks of 10 trees per treatment were sampled on five dates from May to November 1995. Phloem nitrogen and potassium concentrations were elevated in trees on fertilized plots, whereas phloem calcium concentrations were decreased. Fertilization significantly enhanced (10-20%) concentrations of phloem phenolics and proanthocyanidins. In contrast, phloem soluble sugars and starch concentrations were up to 30% lower in fertilized trees than in control trees. Increased phenolic concentrations and lower nonstructural carbohydrates should correlate with reduced tissue palatability and decreased pathogen susceptibility in fertilized trees; however, resin flows were significantly lower (30-100%) in fertilized trees compared with control trees, which may facilitate pine bark beetle establishment. Furthermore, fertilization-induced increases in phloem nitrogen concentration may be more important than tissue carbohydrate or phenolic content in determining tissue palatability.