Allometric relationships predicting foliar biomass and leaf area:sapwood area ratio from tree height in five Costa Rican rain forest species

We developed allometric equations to predict whole-tree leaf area (A(l)), leaf biomass (M(l)) and leaf area to sapwood area ratio (A(l):A(s)) in five rain forest tree species of Costa Rica: Pentaclethra macroloba (Willd.) Kuntze (Fabaceae/Mim), Carapa guianensis Aubl. (Meliaceae), Vochysia ferru-gi-nea Mart. (Vochysiaceae), Virola koshnii Warb. (Myris-ticaceae) and Tetragastris pana-mensis (Engl.) Kuntze (Burseraceae). By destructive analyses (n = 11-14 trees per species), we observed strong nonlinear allometric relationships (r(2) >/= 0.9) for predicting A(l) or M(l) from stem diameters or A(s) measured at breast height. Linear relationships were less accurate. In general, A(l):A(s) at breast height increased linearly with tree height except for Penta-clethra, which showed a negative trend. All species, however, showed increased total A(l) with height. The observation that four of the five species increased in A(l):A(s) with height is consistent with hypotheses about trade--offs between morphological and anatomical adaptations that favor efficient water flow through variation in the amount of leaf area supported by sapwood and those imposed by the need to respond quickly to light gaps in the canopy.     

Changes in whole-tree water relations during ontogeny of Pinus flexilis and Pinus ponderosa in a high-elevation meadow

We measured sap flux in Pinus ponderosa Laws. and Pinus flexilis James trees in a high-elevation meadow in northern Arizona that has been invaded by conifers over the last 150 years. Sap flux and environmental data were collected from July 1 to September 1, 2000, and used to estimate leaf specific transpiration rate (El), canopy conductance (Gc) and whole-plant hydraulic conductance (Kh). Leaf area to sapwood area ratio (LA/SA) increased with increasing tree size in P. flexilis, but decreased with increasing tree size in P. ponderosa. Both Gc and Kh decreased with increasing tree size in P. flexilis, and showed no clear trends with tree size in P. ponderosa. For both species, Gc was lower in the summer dry season than in the summer rainy season, but El did not change between wet and dry summer seasons. Midday water potential (Psi(mid)) did not change across seasons for either species, whereas predawn water potential (Psi(pre)) tracked variation in soil water content across seasons. Pinus flexilis showed greater stomatal response to vapor pressure deficit (VPD) and maintained higher Psi(mid) than P. ponderosa. Both species showed greater sensitivity to VPD at high photosynthetically active radiation (PAR; > 2500 micromol m-2 s-1) than at low PAR (< 2500 micromol m-2 s-1). We conclude that the direction of change in Gc and Kh with increasing tree size differed between co-occurring Pinus species, and was influenced by changes in LA/SA. Whole-tree water use and El were similar between wet and dry summer seasons, possibly because of tight stomatal control over water loss.     

Photosynthesis and water relations of savanna tree species differing in leaf phenology

Godmania macrocarpa Hemsley, a deciduous tree characteristic of fire protected areas of the savanna region of central Venezuela, was more drought tolerant, allocated more N to leaves and had consistently higher photosynthetic rates than Curatella americana L., a ubiquitous species growing successfully within the grasslands of tropical American savannas. Godmania macrocarpa maintained higher leaf conductance and photosynthesized at higher xylem water tensions than C. americana. As the dry season progressed, G. macrocarpa was more affected by water stress than C. americana, which may explain why G. macrocarpa shed its leaves before forming new leaves. For both species, leaf sap osmolality was strongly correlated with, but not completely accounted for by, soluble sugars. Integrated water-use efficiency, as measured by delta(13)C, was similar for both species, but young leaves were more efficient than old leaves. Water-use efficiency of adult leaves was similar in both species as a result of higher photosynthetic rates in G. macrocarpa and lower leaf conductances in C. americana. Compared to G. macrocarpa, instantaneous photosynthetic N-use efficiency was higher in C. americana despite its lower maximum photosynthetic rates. The absence of G. macrocarpa trees from open grasslands, despite their high productive capacity, is possibly the result of unfavorable conditions for germination, poor survival of seedlings, and lack of resistance against fire.     

Influence of drought stress and low irradiance on plant water relations and structural constituents in needles of Pinus ponderosa seedlings

The influence of low light on tolerance to prolonged drought was tested on unshaded and shaded seedlings of ponderosa pine (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Unshaded seedlings of P. ponderosa var. ponderosa were also drought stressed to compare varietal responses to drought. The maximum irradiance received by shaded seedlings was 10% of full light. Seedlings were progressively drought stressed until predawn water potentials (Psi(x)) were -5.0 MPa. Relative water content (RWC) and the reciprocal of Psi(x) were analyzed by means of an unusual application of the pressure-volume relationship for determination of RWC of the apoplast (RWC(a)), osmotic potential at full turgor (Psi(oft)), and ratio of fully turgid weight to dry weight. Major varietal differences in drought response were in RWC(a) and needle cellulose content. The shaded seedlings showed tissue damage at relative water contents < 60%, and were killed by water deficits from which unshaded seedlings recovered. Correspondingly, shaded plants had significantly higher cell volume/cell mass ratio, Psi(oft), less cellulose in needle tissue, and lower RWC(a) than unshaded plants. These differences suggest that low irradiance restricts drought adaptation in ponderosa pine.     

Effect of pruning on branch production and water relations in widely spaced ponderosa pines

This study dealt with the effects of pruning on branch and leaf area (F _a) production of ponderosa pines growing in silvopastoral systems in Patagonia. We hypothesized that pruning positively influences the number of branches per whorl and their basal area growth rate, changing F _a production. In addition, we studied some water relations in order to explain potential differences in branch growth rates between treatments. Two mathematical models were developed to estimate branch and total F _a. The averaged diameter at the third year of pruning was, for high-pruned trees 3.1 and 3.6 cm at the bottom and middle of the crown, against 4 and 4.4 cm for low-pruned trees. Pruning did not produce changes in the number of branches per whorl (approximately 7.6 branches per whorl). Water stress may be responsible of this lower branch growth in pruned trees. Water potential, stomatal conductance and transpiration were lower in high- than in low-pruned trees.     

Above- and belowground responses to tree thinning depend on the treatment of tree debris

Mechanical mastication is increasingly prescribed for wildfire mitigation, yet little is known about the ecological impacts of this fuels treatment. Mastication shreds trees into woodchips as an alternative to tree thinning and burning the resulting slash, which can create soil disturbances that favor exotic plants. Previous research on mastication has not simultaneously considered both the responses of soil organisms and understory plant communities. We compared mastication to slash pile burning (both 6-months and 2.5-years post-treatment) and untreated controls in pinyon–juniper (Pinus edulis–Juniperus osteosperma) woodland and measured soil properties, arbuscular mycorrhizal fungi (AMF) and understory plant composition. Our results showed that slash pile burns had severely degraded soil properties and low AMF abundance and richness compared to untreated or mastication plots. Pile burns were dominated by exotic plant species and had approximately 6× less understory plant abundance and richness than untreated plots. Only two variables differed between mastication and untreated plots 6-months post-treatment: mastication had lower soil temperature and higher soil moisture. Mastication plots 2.5-years post-treatment had more plant cover and richness than untreated plots or pile burns, although non-native Bromus tectorum cover was also greater and AMF spore richness was lower than untreated plots. The structural equation model (SEM) we developed showed that plant cover strongly influenced AMF abundance (0.50) and both plant cover (0.36) and AMF (0.31) positively influenced soil stability. In the short-term, mastication is a preferable method as it creates fewer disturbances than pile burning; however long-term impacts of mastication need further study as this practice could affect native plant communities. Our results suggest that the manner in which woody debris is treated following tree thinning has an important influence on soil stability and native plant biodiversity.     

Dry-season leaf flushing of Enterolobium cyclocarpum (ear-pod tree): above- and below-ground phenology and water relations

Above- and belowground phenology and water relations of Enterolobium cyclocarpum Jacq. trees in the dry forest of Santa Rosa National Park, Costa Rica were studied during two consecutive phenological cycles, from November 1998 to June 2000. Aboveground phenological activity, including leaf shedding, growth and maturation of dormant fruits, new leaf flushing and flowering, occurred during the dry season. Measurements of leaf water potential, stomatal conductance and sap flow indicated that stomata of newly flushed leaves remained essentially closed until the onset of the first rains, suggesting that the main factor accounting for the favorable water balance of dry-season flushed leaves was their capacity to restrict water loss. Evidence of a contribution from stem and root water stores to shoot expansion was mixed because only the first dry-season flushing episode monitored was accompanied by a marked decrease in stem and root water potentials. Fine root production did not precede leaf flushing, occurred only after the onset of the rainy season and stopped under drought conditions, suggesting that soil water content was the most important variable controlling fine root dynamics in this species.     

Influence of precommercial thinning grade on branch diameter and crown ratio in Pinus sylvestris in southern Sweden

Precommercial thinning is regarded as one of the most important measures for influencing timber quality in stands of Scots pine (Pinus sylvestris L.). The present study considers the influence of both thinning intensity and tree height at the time of thinning on diameter of the thickest branch and crown ratio. A total of 115 plots (either naturally regenerated, planted or sown, and either thinned down to 528–8000 stems?ha^?1 or untreated) included in 20 sites in southern Sweden were analysed. The average tree height after thinning varied from 1.2 to 8.3?m. An increasing number of remaining stems resulted in a reduction in branch diameter, although the reduction appeared to be only minor if the number of stems after thinning was more than 3000 stems?ha^?1. It was found that late thinning reduced the diameter of the thickest branch. The crown ratio decreased with stand height, number of stems after thinning and average height at thinning. The results were consistent for all trees and for the 500 thickest trees per hectare.     

Growth of young Pinus ponderosa and Pinus contorta on compacted soil in central Washington

Growth in height, diameter, and volume was measured on 9- to 18-year-old ponderosa pine (Pinus ponderosa) and 10- to 13-year-old lodgepole pine (Pinus contorta) trees growing on or near compacted skid trails in the Yakima Indian Reservation in south-central Washington. Soil bulk density of the 0- to 30.5-cm deep layer was measured with a single-probe nuclear densimeter on two sides of each sample tree and in adjacent undisturbed soil. On three ponderos pine sites logged 23 years before the study, average bulk density on skid trails was 15% greater than on adjacent undisturbed soil. On a lodgepole pine site logged 14 years before the study, soil on skid trails averaged 28% greater bulk density than undisturbed soil.Total growth of ponderosa pine and the last 5 years of growth were significantly related (P = 0.07) by regression analysis to age of trees, site index, basal area of the adjacent overstory, and the percentage of increase in soil bulk density. At the mean increase in soil bulk density, total height, diameter, and volume growth were reduced 5%, 8%, and 20%, respectively.Total growth of lodgepole pine and the last 5 years of height, diameter, and volume growth were significantly related to tree age and the percentage of soil organic matter. Increase in soil bulk density was not significantly related to growth of this species.     

Differences in leaf gas exchange and water relations among species and tree sizes in an Arizona pine-oak forest

We compared leaf gas exchange and water potential among the dominant tree species and major size classes of trees in an upland, pine-oak forest in northern Arizona. The study included old-growth Gambel oak (Quercus gambelii Nutt.), and sapling, pole, and old-growth ponderosa pines (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Old-growth oak had higher predawn leaf water potential (Psi(leaf)) than old-growth pine, indicating greater avoidance of soil water stress by oak. Old-growth oak had higher stomatal conductance (G(w)), net photosynthetic rate (P(n)), and leaf nitrogen concentration, and lower daytime Psi(leaf) than old-growth pine. Stomatal closure started at a daytime Psi(leaf) of about -1.9 MPa for pine, whereas old-growth oak showed no obvious reduction in G(w) at Psi(leaf) values greater than -2.5 MPa. In ponderosa pine, P(n) and G(w) were highly sensitive to seasonal and diurnal variations in vapor pressure deficit (VPD), with similar sensitivity for sapling, pole, and old-growth trees. In contrast, P(n) and G(w) were less sensitive to VPD in Gambel oak than in ponderosa pine, suggesting greater tolerance of oak to atmospheric water stress. Compared with sapling pine, old-growth pine had lower morning and afternoon P(n) and G(w), predawn Psi(leaf), daytime Psi(leaf), and soil-to-leaf hydraulic conductance (K(l)), and higher foliar nitrogen concentration. Pole pine values were intermediate between sapling and old-growth pine values for morning G(w) and daytime Psi(leaf), similar to sapling pine for predawn Psi(leaf), and similar to old-growth pine for morning and afternoon P(n), afternoon G(w), K(l), and foliar nitrogen concentration. For the pines, low predawn Psi(leaf), daytime Psi(leaf), and K(l) were associated with low P(n) and G(w). Our data suggest that hydraulic limitations are important in reducing P(n) in old-growth ponderosa pine in northern Arizona, and indicate greater avoidance of soil water stress and greater tolerance of atmospheric water stress by old-growth Gambel oak than by old-growth ponderosa pine.     

Spatial and temporal variations in leaf area index, specific leaf area and leaf nitrogen of two co-occurring savanna tree species

Foliage growth, mass- and area-based leaf nitrogen concentrations (Nm and N a) and specific leaf area (SLA) were surveyed during a complete vegetation cycle for two co-occurring savanna tree species: Crossopteryx febrifuga (Afzel. ex G. Don) Benth. and Cussonia arborea A. Rich. The study was conducted in the natural reserve of Lamto, Ivory Coast, on isolated and clumped trees. Leaf flush occurred before the beginning of the rainy season. Maximum leaf area index (LAI), computed on a projected canopy basis for individual trees, was similar (mean of about 4) for both species. Seasonal courses of the ratio of actual to maximum LAI were similar for individuals of the same species, but differed between species. For C. febrifuga, clumped trees reached their maximum LAI before isolated trees. The LAI of C. arborea trees did not differ between clumped and isolated individuals, but maximum LAI was reached about 2 months later than for C. febrifuga. Leaf fall was associated with decreasing soil water content for C. arborea. For C. febrifuga, leaf fall started before the end of the rainy period and was independent of changes in soil water content. These features lead to a partial niche separation in time for light resource acquisition between the two species. Although Nm, N a and SLA decreased with time, SLA and N a decreased later in the vegetation cycle for C. arborea than for C. febrifuga. For both species, N a decreased and SLA increased with decreasing leaf irradiance within the canopy, although effects of light on leaf characteristics did not differ between isolated and clumped trees. Given relationships between N a and photosynthetic capacities previously reported for these species, our results show that C. arborea exhibits higher photosynthetic capacity than C. febrifuga during most of the vegetation cycle and at all irradiances.     

Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus petraea (Matt.) Liebl.)

To quantify the effects of crown thinning on the water balance and growth of the stand and to analyze the ecophysiological modifications induced by canopy opening on individual tree water relations, we conducted a thinning experiment in a 43-year-old Quercus petraea stand by removing trees from the upper canopy level. Soil water content, rainfall interception, sap flow, leaf water potential and stomatal conductance were monitored for two seasons following thinning. Seasonal time courses of leaf area index (LAI) and girth increment were also measured. Predawn leaf water potential was significantly higher in trees in the thinned stand than in the closed stand, as a consequence of higher relative extractable water in the soil. The improvement in water availability in the thinned stand resulted from decreases in both interception and transpiration. From Year 1 to Year 2, an increase in transpiration was observed in the thinned stand without any modification in LAI, whereas changes in transpiration in the closed stand were accompanied by variations in LAI. The different behaviors of the closed and open canopies were interpreted in terms of coupling to the atmosphere. Thinning increased inter-tree variability in sap flow density, which was closely related to a leaf area competition index. Stomatal conductance varied little inside the crown and differences in stomatal conductance between the treatments appeared only during a water shortage and affected mainly the closed stand. Thinning enhanced tree growth as a result of a longer growing period due to the absence of summer drought and higher rates of growth. Suppressed and dominant trees benefited more from thinning than trees in the codominant classes.     

Growth and water relations of Eucalyptus marginata (jarrah) stands in response to thinning and fertilization

We studied the effects of five thinning treatments (T1 = 5.5, T2 = 11, T3 = 16.5, T4 = 22.5 and T5 = 28.5 m(2) ha(-1) basal area under bark) x two fertilizer treatments (F0 = unfertilized and F1 = fertilized with 400 kg ha(-1) N plus 229 kg ha(-1) P) on growth and water relations of pole-sized Eucalyptus marginata J. Donn ex Sm. trees growing in southwestern Australia. Thinning reduced leaf area index (LAI) from 2.1 in the T4 and T5 treatments to 0.8 in the T1F0 treatment. Fertilizer had no effect on LAI in the T2, T4 or T5 treatments, but increased LAI by 45 and 20% in the T1 and T3 treatments, respectively. Thinning plus fertilizing increased diameter growth most in the fastest growing trees, from 0.4 cm year(-1) for trees in the T5F0 and T5F1 treatments to 0.7 and 1.2 cm year(-1) for trees in the T1F0 and T1F1 treatments, respectively. In both fertilizer treatments, stand basal area and volume growth increased with increasing stand density up to 15 m(2) ha(-1), and thereafter declined with increasing stand density, such that the growth rate of trees in the T5 treatment was only half of that at a stand density of 15 m(2) ha(-1). In response to fertilizer, growth rates of the slowest and fastest-growing trees increased from 0.35 and 3.5 m(2) ha(-1) year(-1) (F0) to 0.56 and 5.4 m(3) ha(-1) year(-1) (F1), respectively. Stand growth efficiency (growth per unit LAI) increased in response to thinning, and fertilizer increased stand growth efficiency at all stand densities. Throughout the dry season, T5 trees had lower predawn shoot water potentials (Psi(pd)) (minimum of -1.5 MPa) than T1 or T2 trees (minimum of -0.7 MPa). Fertilizer decreased Psi(pd) in T5 trees (by -0.9 and -1.5 MPa, respectively, in F0 and F1), but not in T1 or T2 trees. Stand growth rate was closely related to cumulative midday water stress (CMWS) over the dry season, and volume growth rate declined sharply from 6 m(3) ha(-1) year(-1) at a CMWS of 130 MPa days, to zero at a CMWS of 220 MPa days. Application of fertilizer to thinned stands increased LAI, stand growth efficiency and stand growth. In unthinned stands, fertilizer increased stand growth efficiency and stand growth; however, it also increased tree water stress, which limited the fertilizer-induced increases in LAI and growth. We attribute the increase in tree and stand growth in response to application of fertilizer to increased photosynthetic rates, increased allocation to stem wood, and in thinned stands also to higher LAIs.     

Effect of thinning intensity on tree growth and temporal variation of seed and cone production in a Pinus koraiensis plantation

Thinning of Korean pine (Pinus koraiensis Sieb. et Zucc.) is used to facilitate timber and cone production. The present study in Northeast China investigated the effects of thinning intensity on individual tree growth, temporal variation in cone yield, and seed quality in Korean pine plantation. In 2005, five thinning intensity levels (none, extreme, heavy, moderate and light) were set in 15 permanent plots in a 32-year-old Korean pine plantation at Mengjiagang Forest Farm, Jiamusi City, Heilongjiang Province. We recorded tree growth and seed cone production from 2013 to 2016, i.e., from 8 to 11 years after thinning. Except for height growth, thinning increased tree growth (diameter at breast height and crown size) and improved cone yield. The extreme thinning treatment (to 300 trees per hectare) resulted in the largest tree diameter, tree volume, crown size and 4-year cone production per tree. The highest cone yield per tree in the mast year (2014) was observed when stands were thinned to 500 trees per hectare (heavy thinning). Although the best cone and seed quality and the largest cone and seed mass per tree were recorded in the heavily thinned stand, no significant differences were found between heavy and moderate thinning stands (750 trees per hectare). At the stand level, the moderately thinned stand had the highest basal area, stock volume and seed cone production per stand. Our results suggest that thinning to 750 trees per hectare will improve timber and cone productivity in 40-year-old P. koraiensis stands.     

Responses of apple fruit size to tree water status and crop load

The combined effects of irrigation rate and crop load on apple yield and fruit size were examined in two commercial apple orchards (cv. Golden Delicious) in a semi-arid zone. The irrigation rates applied were 1, 3 and 7 mm day(-1), and the two fruit thinning treatments involved adjusting crop load to 100 and 300 fruits per tree at Ortal and 50 and 150 fruits per tree at Matityahu. Unthinned trees served as the control. The fruit from each tree was picked separately, and fruit size distribution was determined with a commercial grading machine. Midday stem water potentials varied from -0.9 to -2.8 MPa, crop load varied from 80,000 to 1,900,000 fruit ha(-1) and crop yield varied from 10 to 144 Mg ha(-1). Midday stem water potential decreased with increasing crop load in all irrigation treatments at Matityahu, but only in the 1 mm day(-1) treatment at Ortal. The extent of the lowering of midday stem water potential by crop load decreased with increasing soil water availability. At both orchards, a similar response of total crop yield to crop load on a per hectare basis was observed. Mean fruit mass and relative yield of fruit > 70 mm in diameter increased with midday stem water potential, with the low crop loads having similar but steeper slopes than the high crop load. The responses of mean fruit mass and relative yield of fruit > 70 mm in diameter to midday stem water potential were similar at both orchards, perhaps indicating that thresholds for irrigation scheduling are transferable to other orchards within a region. Factors that may limit the transferability of these thresholds are discussed.