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.     

Water relations of pine seedlings in contrasting overstory environments

Overstory conditions influence understory microclimate and resource availability, leading to gradients in evaporative demand and moisture availability that influence seedling water relations. Partial canopies may either reduce seedling moisture stress by ameliorating environmental conditions, or increase moisture stress by reducing soil moisture availability. This study used stable isotope ratios of oxygen (δ¹⁸O) and carbon (δ¹³C) and mass-based foliar nitrogen concentrations to investigate changes in transpiration (E), stomatal conductance (g_s) and intrinsic water use efficiency (iWUE) of pine seedlings across an overstory gradient from open canopy gap environments to closed canopy forest. Foliar δ¹⁸O increased sharply from basal areas of 0–10 m² ha⁻¹ in Pinus banksiana, Pinus resinosa, and Pinus strobus seedlings, followed by a more gradual increase with further increases in basal area. Foliar δ¹³C followed a similar, but less pronounced pattern in P. banksiana and P. strobus seedlings, and had no apparent relationship with overstory basal area in P. resinosa seedlings. The slope of the δ¹⁸O:δ¹³C relationship was positive for every species. Foliar nitrogen concentrations were not correlated with overstory basal area. These results suggest seedling E declined as overstory basal area increased due to reductions in g_s, while iWUE increased slightly from open gaps to partial canopy environments. Open gap environments appear to provide sufficient moisture to sustain high leaf-level gas exchange rates in the species we studied, while relatively small increases in overstory basal area apparently promote rapid declines in g_s, leading to greatly reduced seedling water loss and small increases in iWUE.     

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.     

Performance of loblolly pine (Pinus taeda L.) seedlings and micropropagated plantlets on an east Texas site: II. Water relations

Loblolly pine (Pinus taeda L.) seedlings and plantlets, produced from drought resistant genetic families were planted in east Texas and their relative water relations compared through the first two, sixth and seventh growing seasons. Stock type (seedling or plantlet) had a larger influence on water relations than did family source in the early years. Seedlings exhibited more of those characteristics generally considered to benefit survival under harsh environments. The principal difference between stock types was the degree of progressive decrease in predawn water potential (Ψ_pre) through the growing season. On average and across families, Ψ_pre decreased through the first growing season by 32 and 17% for plantlets and seedlings, respectively.Complete stomatal closure was never observed for any genetic source or stock type even when water potential was below −2.0 MPa. However, seedlings exhibited more stomatal regulation by lowering their stomatal conductance when water deficit was encountered, and increasing it during periods of favorable soil moisture availability. Predawn water potential was never below −0.9 MPa for any tree in either of the two first growing seasons. Although significant differences between seedling and plantlet performance were observed, their magnitude was small and diminished in the later years (sixth and seventh growing seasons). We conclude that plantlets need further development to be considered as a viable regeneration source for loblolly pine plantations on sites where summer drought is common.     

Water use in thinned loblolly pine plantations

Soil moisture and throughfall were monitored for two growing seasons under three thinning intensities: basal areas of 7.8 m²ha⁻¹, 12.6 m²ha⁻¹ and 26.6 m²ha⁻¹ (unthinned control) in an 11- to 12-year-old loblolly pine (Pinus taeda L.) plantation. The first year after thinning soil water decreased at a rate of 0.8 mm day⁻¹, 1.0 mm day⁻¹ and 1.4 mm day⁻¹, respectively, in the 7.8 m²ha⁻¹, 12.6 m²ha⁻¹, and 26.6 m²ha⁻¹ basal area plots between May and late August. The more rapid development of soil moisture deficits in the unthinned stand was owing to both greater soil moisture use and reduced throughfall. For the same time period in the second year, with below-normal rainfall, soil moisture deficits developed at a rate of 1.5, 1.5 and 1.7 mm day⁻¹ for the two heavily thinned and the unthinned treatments. Estimated growing season soil water use rates were 4.4 mm day⁻¹, 4.2 mm day⁻¹, and 4.0 mm day⁻¹, respectively for the 7.8 m²ha⁻¹, 12.6 m²ha⁻¹ and 26.6 m²ha⁻¹ basal area levels in Year 1. The second year after thinning water use rates for the same period were 2.7 mm day⁻¹, 2.6 mm day⁻¹, and 2.5 mm day⁻¹, respectively. The results of this study suggest that the capacity to manage available water in loblolly pine stands by thinning is more a function of reduced interception loss and increased throughfall than reduced water use and is also weather dependent. Growing seasons with low rainfall or only one or two large rainfall events will not permit much opportunity to manage soil moisture.     

Seasonal changes in intensity of bud dormancy in loblolly pine seedlings

The terminal buds of six-month-old loblolly pine (Pinus taeda L.) seedlings remained closed for approximately six months, although bud dormancy, as measured by rate of bud break in a standard greenhouse environment, was only exhibited for about one month. The peak of bud dormancy was in December for seedlings grown near Auburn, Alabama. However, the timing and intensity varied with seed source and may have been affected by the warm fall temperatures. Seedlings from the more northern provenances entered dormancy first and reached a deeper state of dormancy than seedlings from southern provenances. The rate of shoot elongation was not consistently related to the rate of bud break.     

Early gains from planting large-diameter seedlings and intensive management are additive for loblolly pine

A seedling size/intensive management study with Pinustaeda L. was established in 1993 on two sites in the CoastalPlain of Georgia and South Carolina. Each site contained a 2 × 2split-plot study involving two seedling sizes and two levels ofestablishment intensity. Ideotype ``B' seedlings averaged 5.0 mmin diameter (at the root collar) and were 43 cm tall. Ideotype``A' seedlings averaged 8.5 mm in diameter and were 50 cm tall. ``Standard' establishment practices included herbicides(hexazinone and sulfometuron) and fertilizer (DAP) appliedduring the first year. The ``intensive' management involved twoherbicide applications during the first year and two during thesecond year, fertilizer during the first and third years, andinsecticide applications during the first two years (for controlof tip moth, Rhyacionia frustrana Comstock).Intensive management did not affect survival but planting largerseedlings increased survival slightly on one site. However,treatments affected early growth at both sites. On both sites,fourth-year plot-volumes were increased with greaterestablishment intensity and larger seedlings but there was nointeraction between stock size and establishment intensity. Early growth gains were greatest when both intensive managementand larger seedlings were combined. Depending on site, thiscombination resulted in 21% to 51% more volume (at age 4) thanthe next best treatment (standard seedlings with intensivemanagement).     

Long-term prescribed burning increases nutrient uptake and growth of loblolly pine seedlings

Loblolly pine (Pinus taeda L.) seedlings were grown from seed in a greenhouse on A1 horizon soil collected from field plots that have been burned each winter or maintained in an unburned condition for 33 years. Soils from burned and unburned plots were treated with phosphorus (P), P and calcium (Ca), or left untreated. After 32 weeks, height, biomass, and nitrogen (N) and P uptake were greater on soil from burned versus unburned plots, although application of P masked these effects. Addition of P increased plant biomass, seedling height, and uptake of N and P, but depressed levels of soil NH₄N. These results suggest that long-term prescribed burning may have a positive effect on nutrient availability that will benefit seedlings of the next stand.     

Acclimation of loblolly pine (Pinus taeda) seedlings to high temperatures

To determine the extent to which loblolly pine seedlings (Pinus taeda L.) acclimate to high temperatures, seedlings from three provenances-southeastern Texas (mean annual temperature 20.3 degrees C), southwestern Arkansas (mean annual temperature 16.2 degrees C) and Chesapeake, Maryland (mean annual temperature 12.8 degrees C)-were grown at constant temperatures of 25, 30, 35 or 40 degrees C in growth chambers. After two months, only 14% of the seedlings in the 40 degrees C treatment survived, so the treatment was dropped from the experiment. Provenance and family differences were not significant for most measured variables. Total biomass was similar in the 25 and 30 degrees C treatments, and less in the 35 degrees C treatment. Foliage biomass was higher, and root biomass lower, in the 30 degrees C treatment compared with the 25 degrees C treatment. Net photosynthesis and dark respiration of all seedlings were measured at 25, 30 and 35 degrees C. Both net photosynthesis and dark respiration exhibited acclimation to the temperature at which the seedlings were grown. For each temperature treatment, the highest rate of net photosynthesis was measured at the growth temperature. Dark respiration rates increased with increasing measurement temperature, but the basal rate of respiration, measured at 25 degrees C, decreased from 0.617 &mgr;mol m(-2) s(-1) in the 25 degrees C treatment to 0.348 &mgr;mol m(-2) s(-1) in the 35 degrees C treatment, resulting in less carbon loss in the higher temperature treatments than if the seedlings had not acclimated to the growth conditions. Temperature acclimation, particularly of dark respiration, may explain why total biomass of seedlings grown at 30 degrees C was similar to that of seedlings grown at 25 degrees C.     

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.     

Effect of elevated carbon dioxide concentration and root restriction on net photosynthesis, water relations and foliar carbohydrate status of loblolly pine seedlings

To determine the effects of CO(2)-enriched air and root restriction on photosynthetic capacity, we measured net photosynthetic rates of 1-year-old loblolly pine seedlings grown in 0.6-, 3.8- or 18.9-liter pots in ambient (360 micro mol mol(-1)) or 2x ambient CO(2) (720 micro mol mol(-1)) concentration for 23 weeks. We also measured needle carbohydrate concentration and water relations to determine whether feedback inhibition or water stress was responsible for any decreases in net photosynthesis. Across all treatments, carbon dioxide enrichment increased net photosynthesis by approximately 60 to 70%. Net photosynthetic rates of seedlings in the smallest pots decreased over time with the reduction occurring first in the ambient CO(2) treatment and then in the 2x ambient CO(2) treatment. Needle starch concentrations of seedlings grown in the smallest pots were two to three times greater in the 2x ambient CO(2) treatment than in the ambient CO(2) treatment, but decreased net photosynthesis was not associated with increased starch or sugar concentrations. The reduction in net photosynthesis of seedlings in small pots was correlated with decreased needle water potentials, indicating that seedlings in the small pots had restricted root systems and were unable to supply sufficient water to the shoots. We conclude that the decrease in net photosynthesis of seedlings in small pots was not the result of CO(2) enrichment or an accumulation of carbohydrates causing feedback inhibition, but was caused by water stress.     

Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.     

Morphological and physiological differences in Scots pine seedlings of six seed origins

One-year-old seedlings of Scots pine (Pinus sylvestris L.) offour native seed origins (Loch Maree Islands, Glengarry/GlenMorriston, Glen Affric and Abernethy), a commercial Britishseedlot, and a seedlot from Hedesunda, in middle Sweden, werecompared at monthly intervals from October 1993 to April 1994.Seedling morphology, root condition, root frost hardiness andbud dry matter were determined at each date. There were clear morphological differences among seed origins.Seedlings raised from the commercial seedlot (A70) were largerbut had a poorer root:shoot ratio than the other seed origins.Of the native pines tested, the Loch Maree Islands origin allocateda larger proportion of its photosynthate to fine roots and needlesand smaller proportion to woody structures. Seedlings raised from the commercial British seedlot tendedto have poorer bud lignification than the other origins andalso, in autumn, higher electrolyte leakage rates from its fineroots. During winter, the Swedish origin had the lowest fineroot electrolyte leakage. Seedlings of all origins showed aprogressive increase in fine root hardiness towards mid-winterwith maximum hardiness (–7°C) in January. Dehardeningoccurred over subsequent months reaching –3°C in April.Differences among origins were evident. The Swedish seedlotdeveloped greater frost resistance than the other origins, hardeningbegan earlier in autumn and dehardening began later in spring.The commercial seedlot hardened later than the other originsbut reached a similar level of frost hardiness by January. Ofthe native pines, seedlings of the Loch Maree Islands originwere slowest to develop root hardiness.     

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.     

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.     

Growth and water potential of j-rooted loblolly and eastern white pine seedlings over three growing seasons

The effects of j-rooting on water stress and growth of loblolly (Pious taeda L.) and eastern white pine (Pious strobus L.) were examined over three growing seasons in the field. Seedlings were planted in an area with severe herbaceous competition with either their roots planted straight or bent into a j shape. All seedlings were planted with their root collars placed at the soil surface. During the first year j-rooted seedlings consistently had lower water potentials but never statistically significant. Since both treatments were planted with the root collar at the soil surface, this trend was likely due to an initial shallower root system in j-rooted seedlings. In year three no differences in water potential were significant and no trends were evident. Growth did not differ significantly by treatment at any time but, by year three, j-rooted plants were consistently larger for both species.     

Financial performance of diverse levels of early competition suppression and pre-commercial thinning on loblolly pine stand development

New Forests - Competition suppression within loblolly pine plantations (Pinus taeda L.) is typically carried out within the first 2 years of a plantation, but competition control for longer...     

Movement of respiratory CO(2) in stems of loblolly pine (Pinus taeda L.) seedlings

Temperature-independent fluctuations in stem CO(2) efflux were measured in Pinus taeda L. seedlings. Stem CO(2) efflux was measured during high and low transpiration rates, high and low net photosynthesis rates, and normal and interrupted substrate supply conditions. Stem CO(2) efflux rates were an average of 6.7% lower during periods of high transpiration compared to periods of low transpiration. This difference in stem CO(2) efflux rates was not due to water stress. The most likely cause was movement of respiratory CO(2) in the transpiration stream. Interruption of substrate supply to the stem by phloem girdling reduced stem CO(2) efflux rates. Increasing net photosynthesis rates from low to high had no effect on stem CO(2) efflux, but decreasing net photosynthesis from high to low caused relatively small reductions in stem CO(2) efflux. These results indicate that diurnal changes in net photosynthesis rate may play a small role in temperature-independent afternoon depressions of stem CO(2) efflux. The transport of respiratory CO(2) by the transpiration stream compromises measurements of woody tissue respiration obtained by commonly accepted gas exchange techniques. This phenomenon could also affect measurement of leaf net photosynthesis and branch woody tissue respiration.     

Durability of selected mulches,their ability to control weeds,and influence growth of loblolly pine seedlings

Several mulches of natural, synthetic, or blends of natural and synthetic fibers were tested around newly planted loblolly pine (Pinus taeda L.) seedlings on a sheared and windrowed site in central Louisiana, USA. The vegetation was primarily winter annuals, some residual grasses and forbs, and sparse woody regrowth. Study 1 was rotary mowed just prior to planting in March 1992, and 35 mulches and an untreated check were established. In Study 2, 15 mulches and an untreated check were established in a 1-year-old rough in March 1993. In both studies, a single loblolly pine seedling formed each plot established in a randomized complete block design, with 10 blocks as replicates. Each block was planted with a separate open-pollinated loblolly pine family.Nearly all mulches had deteriorated to some extent after three growing seasons. Synthetic mulches were generally more durable than the natural or natural/synthetic mulches. Mulching eliminated the established vegetation and germinants, and vegetation did not readily reestablish following the deterioration of a mulch. The soil seed bank apparently was not sufficient to regenerate areas that were once covered with mulch and many of the natural materials deteriorated into a fibrous cover that acted like a natural litter layer. Both of these residual weed control effects -- insufficient soil seed bank and formation of a fibrous cover -- were important in stopping vegetation from reestablishing after a mulch had deteriorated. After three growing seasons, the loblolly pine seedlings generally grew better if mulches were used.     

Water relations and photosynthesis in three families of radiata pine seedlings known to differ in their response to weed control

Needle water potential (Ψ_n), stomatal resistance (r_s) and net photosynthesis (P_N) were measured in potted radiata pine seedlings from three families (A, B, and C) during a soil drying and rewritting event. Measurements were made in a controlled environment of 25°C, 14-h days and 18°C, 10-h nights; vapour pressure deficit of 16 mbar; photon flux of 400 μE m^?2 s_?1; and wind speed of 1 m s^?1.Stomatal resistance increased at a greater rate in family A than in family B than in family C as Ψ_n decreased during the drying phase: at ?25 bar Ψ_n, r_s values were 77, 60, and 46 s cm^?1 for families A, B and C respectively. Net photosynthesis (P_N) was greatest in family A at high Ψ_n but least at low Ψ_n: at Ψ_n ?9 bar, P_N was 9.4 6.4, and 8.2 ng cm^?2 s^?1 for families A, B, and C respectively, but for ?25 bar Ψ_n corresponding P_N values were 0.4, 1.1, and 0.9.Consequently family A had a greater proportional decrease in both P_N and water use efficiency from high Ψ_n to low Ψ_n. This may help explain in part observed field behaviour (P.P. Cotterill, unpublished) where family A produces well relative to other families when weeds are controlled (high Ψ_n condition) but produces less well relative to other families when weeds are not controlled (low Ψ_n condition).Recovery of Ψ_n after rewatering preceded that of r_s and P_N, and recovery of P_n was typically biphasic with 70% of recovery coinciding with recovery in Ψ_n and the remaining 30% more closely coinciding with recovery in r_s. Recovery was slower in family A than the other two families.