Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Controls over leaf and litter calcium concentrations among temperate trees

Four-fold variation in leaf-litter Ca concentration among 14 tree species growing in a common garden in central Poland was linked to variation in soil pH, exchangeable Ca, soil base saturation, forest floor turnover rates, and earthworm abundance. Given the potential importance of tissue Ca to biogeochemical processes, in this study we investigated potential controls on leaf Ca concentrations using studies of both laboratory seedlings and 30-year-old trees in the field. We first assessed whether species differences in Ca concentration of green leaves and leaf litter were due to differences in Ca uptake, plant growth, or Ca translocation to different organs, by measuring seedlings of 6 of the 14 species grown under controlled conditions of varying Ca supply. We also investigated whether trees species with high Ca concentrations in green leaves and leaf litter access soil Ca to a greater extent than low-Ca species by growing more fine roots in high-Ca soil horizons. Root distribution in the field was determined in all 14 tree species by profile wall mapping and soil sampling of excavated pits. There was no correlation between horizon root count density (number of roots m⁻²) and exchangeable soil Ca, nor was there a correlation of stand-level leaf litter Ca with density of roots 45–100 cm deep in the soil, suggesting that a deeper root distribution does not result in greater Ca acquisition among these species. Variation among species in leaf Ca concentration of greenhouse seedlings was positively correlated with leaf Ca concentrations of mature trees, indicating that the same ranking in leaf Ca among species existed under controlled Ca supply. Species also differed in seedling growth response to Ca supply. Tilia, the species with the highest leaf Ca in the field, generated only 10% as much biomass and height at low relative to high Ca supply, whereas the other species exhibited no significant differences. Species exhibited differences in (i) partitioning of whole plant Ca and biomass to leaf, stem and root organs and (ii) the pattern of such partitioning between high and low Ca treatments. Our data support the hypothesis that although soil Ca supply can contribute to variation among trees in leaf and litter Ca concentration, innate physiological differences among species also can be a major cause for species variation.     

Organic phosphorus in soil water under a European beech (Fagus sylvatica L.) stand in northeastern Bavaria, Germany: seasonal variability and changes with soil depth

Organically bound phosphorus (P) is a mobile form of phosphorus in many soils and thus its dynamics relevant for the leaching and cycling of this element. Despite its importance, little is known about the chemical composition of dissolved organic P. We studied the concentrations, fluxes, and chemical composition of organic P in forest floor leachates and soil solutions in a Rendzic Leptosol under a 90-year-old European beech (Fagus sylvatica L.) forest over a 27-month period (1997–1999). The chemical composition of organic P was analysed using XAD-8 fractionation and ³¹P-nuclear magnetic resonance (NMR) spectroscopy. Organic P was the dominant P form in forest floor leachates as well as in porewaters of the mineral soil. The largest concentrations of organic P were observed during summer and peaked (330–400 g dissolved organic P l^–1) after rain storms following short dry periods, concurrently with the concentrations of organic carbon (OC). Because of high rainfall, fluxes of organic P (and C) were greatest in autumn although concentrations of organic C and P were lower than in summer. In forest floor leachates, the hydrophilic fraction of dissolved organic matter contained 83 ± 13% of the bulk organic P. In soil solutions from 90 cm depth, organic P was almost exclusively in the hydrophilic fraction. Because of the low retention of the hydrophilic fraction of dissolved organic matter in the mineral soils, concentrations of organic P in soil water remained almost constant with depth. Consequently, organic P contributed > 95% of the total P leached into deeper subsoils. The overall retention of organic P in the weakly developed mineral soils was little and so the average annual fluxes of organic P in subsoils at 90 cm depth (38 mg m^–2) comprised 67% of those from the forest floors (57 mg m^–2) during the study period. Hence, organic P proved to be mobile in the studied soil. ³¹P-NMR spectroscopy confirmed the dominance of organic P species in soil water. Signals due to inorganic P occurred only in spectra of samples collected in winter and spring months. Spectra of samples from summer and autumn revealed traces of condensed phosphates. Due to low P contents, identification of organic P species in samples from winter and spring was not always possible. In summer and autumn, monoester and diester phosphates were the dominant organic species and varied little in their relative distributions. The distribution of organic species changed little from forest floor leachates to the subsoil solutions indicating that the composition of P-containing compounds was not influenced by sorptive interactions or biological transformation.     

Leaf litter decomposition and nutrient dynamics in a subtropical forest after typhoon disturbance

Decomposition of typhoon-generated and normal leaf litter and their release patterns for eight nutrient elements were investigated over 3 yr using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. Two common tree species, Castanopsis sieboldii and Schima wallichii, representative of the vegetation and differing in their foliar traits, were selected. The elements analyzed were N, P, K, Ca, Mg, Na, Al, Fe and Mn. Dry mass loss at the end of study varied in the order: typhoon green leaves > typhoon yellow leaves > normal leaves falling for both species. For the same litter type, Schima decomposed faster than Castanopsis. Dry mass remaining after 2 yr of decomposition was positively correlated with initial C:N and C:P ratios. There was a wide range in patterns of nutrient concentration, from a net accumulation to a rapid loss in decomposition. Leaf litter generated by typhoons decomposed more rapidly than did the normal litter, with rapid losses for N and P. Analysis of initial quality for the different litter types showed that the C:P ratios were extremely high (range 896 – 2467) but the P:N ratios were < 0.05 (range 0.02 – 0.04), indicating a likely P-limitation for this forest. On average 32% less N and 60% less P was retranslocated from the typhoon-generated green leaves than from the normal litter for the two species, Castanopsis and Schima. An estimated 2.13 g m^–2 yr^–1 more N and 0.07 g m^–2 yr^–1 more P was transferred to the soil as result of typhoon disturbances, which were as high as 52% of N and 74% of P inputted from leaf litter annually in a normal year. Typhoon-driven maintenance of rapid P cycling appears to be an important mechanism by which growth of this Okinawan subtropical forest is maintained.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

Oak trees and soil interactions in Mediterranean forests: a positive feedback model

Questions: What is the spectrum of variability of chemical elements in a Mediterranean forest ecosystem across the different compartments? Do co‐existing tree species with different leaf chemical composition and nutrient cycling distinctly modify soil conditions? Could these species‐specific, tree‐generated soil changes create a potential positive feedback by affecting long‐term species distribution? Location: Mixed oak forests of southern Spain, Los Alcornocales Natural Park. Methods: We sampled and chemically analysed five different ecosystem components: leaves, leaf fall, litter and superficial (0–25 cm) and sub‐superficial (25–50 cm) soil beneath the canopies of evergreen Quercus suber and deciduous Q. canariensis trees. We used multiple co‐inertia analysis (MCoA) to conjointly analyse the patterns of variability and covariation of eight macro‐ and micronutrients determined in each of the sampled ecological materials. We implemented a path analysis to investigate alternative causal models of relationships among the chemical properties of the different ecosystem components. Results: Variability in the concentration of chemical elements was related to the nature of their biogeochemical cycles. However, the rank of element concentration was consistent across ecosystem components. Analysis of co‐inertia (MCoA) revealed that there was a common underlying multivariate pattern of nutrient enrichment in the ecosystem, which supported the hypothesis of a separation in biogeochemical niches between the two co‐existing oak species, with Q. canariensis having richer plant tissues and more fertile soil directly under each tree than Q. suber. The feasibility of a potential tree–soil positive feedback model was the only statistically validated among several alternative (non‐feedback) models tested. Conclusions: In the studied Mediterranean forests, oak species distinctly modify soil fertility conditions through different nutrient return pathways. Further investigation is needed to address whether these tree‐generated soil changes could affect seedling establishment and ultimately influence species distribution.     

Soil pH accounts for differences in species distribution and leaf nutrient concentrations of Brazilian woodland savannah and seasonally dry forest species

Understanding the environmental factors shaping savannah and tropical forest boundaries is important to predict tropical vegetation responses to climate change and other human-mediated disturbances. To better understand the soil characteristics affecting the distribution of Cerradão (Brazilian woodland savannah) and seasonally dry forest (SDF), two vegetation types occurring next to each other in a similar seasonal climate in south-eastern Brazil, we compared several leaf chemical and morphological traits associated with soil pH and resource availability of Cerradão and SDF woody species. Leaf functional traits were measured for 25 Cerradão and 27 SDF species. We performed between-site comparisons with either all species pooled using phylogenetically independent contrasts or species shared between Cerradão and SDF, as well as congeneric pairs. We found higher specific leaf area and leaf nitrogen, potassium, calcium and sulfur concentrations for SDF species. We did not find higher concentrations for leaf phosphorus and manganese (Mn) for SDF species, despite a higher concentration of these nutrients in SDF soil. Cerradão plants had higher leaf iron (Fe), Mn and aluminium (Al) concentrations. For most of the traits assessed, variance was higher among species and genera than between sites. Nutrients with greater availability in the SDF soil did not invariably exhibit higher concentrations in the leaves of SDF species, indicating that these were not limiting for plant productivity. Higher concentrations in the leaves of Cerradão species for Al, Fe and Mn are probably a consequence of lower soil pH, which increases the availability of these elements. In spite of the differences in belowground resources and the divergence for some traits between Cerradão and SDF, our results show high diversity in leaf functional traits within communities and a similarity of leaf functional traits in closely related species in the contrasting habitats. Besides, we surmise that soil pH is an important factor affecting Cerradão and SDF species distribution, excluding SDF species from more acidic soils, because of the toxic effects of Al, and possibly also Mn and Fe.     

Leaf thickness and UV-B absorbing pigments of plants in relation to an elevational gradient along the Blue Mountains,Jamaica

Terrestrial plant species vary widely in their adaptation to (increasing) solar UV-B radiation. Among the various responses of higher plants to enhanced UV-B are increasing leaf thickness and increasing concentrations of UV-B absorbing compounds. In some (UV-B resistant) plant species increased leaf thickness and UV-B absorbance may form part of mechanisms protecting plants from UV-B damage. However, in UV-B sensitive plant species leaf thickness and UV-B absorbance may increase as well with enhanced UV-B radiation. In the latter case however, this response cannot prevent plant damage and disturbance. In the present field study the relationship between these plant parameters and a natural elevational UV-B gradient on the tropical island of Jamaica was described. Four plant species of the Blue Mountain Tropical Montane Forest, occurring on open forest sites along the roadside and paths were studied along an elevational gradient. Plant species studied are Redbush (Polygonum chinense), Wild ginger (Hedychium gardneranum), John Crow Bush (Bocconia frutescens) and White clover (Trifolium repens). The elevational sites were at 800, 1000, 1200, 1400 and 1600 m above sea level. Leaf thickness was measured of leaves of intact plants around midday in the field. Leaf disks (5 mm) were sampled and extracted with a methanol/HCl mixture. UV-B absorption of these leaf extracts was measured spectrophotometrically. For all species leaves from higher elevations were thicker than those from lower elevations. In addition, the absorption of UV-B of leaf extracts increased with increasing elevations. It is assumed that the calculated gradient of the UV-B_BE from 800 m above sea level: 9.45 kJ m^-2 day^-1 to 9.75 kJ m^-2 day^-1 at 1600 m is related to the measured increase of leaf thickness and UV-B absorbing compounds. The responsiveness of these plant parameters to the elevational gradient does not necessarily imply that the plant species are UV-B resistant. One possibility is that the species studied, which are growing on open, disturbed sites on the forest floor and along mountain-roads, are relatively sensitive to UV-B. In addition to clear sky conditions, mist and clouds occur frequently in this tropical mountane forest at Jamaica. Also, the low nutrient status of the soil (low pH, nutrient deficiency) and the high content of polyphenols in leaves of many plant species of the tropical montane rain forest may relate to the marked response of the species studied with increasing elevation. Abbreviations: asl – above sealevel, UV-B – ultraviolet-B radiation (280–320 nm), TMCF – Tropical Montane Cloud Forest.     

Litterfall and nutrient dynamics in a montane moist evergreen broad-leaved forest in Ailao Mountains,SW China

Montane moist evergreen broad-leaved forest, dominated byLithocarpus and Castanopsis species,is the most extensive stand of subtropical mountain in Yunnan Province, SWChina. Litter production, standing crop of litter on forest floor and nutrientreturn patterns were studied over nine years (1991–1999) in a stand ofprimary evergreen broad-leaved forest in northern crest of the Ailao MountainRange. There were significant yearly variations in litter production, which ismainly related with the masting year of canopy species, and exceptionalphysicalevents (strong winds and snow) in the natural forest. The mean annual smalllitterfall is 7.12 t ha^–1 yr^–1ofwhich leaf litter account for 65% of the total litterfall. The seasonality ofsmall litterfall was bia-modal, with the main one in the late dry season(April–May) and a lesser one in early winter (October–November).Decomposition quotient value was relatively low with 0.58 for total smalllitterfall. Mean large-wood ( 2.5 cm in diameter) ranged from0.21 to 1.41 t ha^–1 yr^–1 with amean of 0.52 t ha^–1 yr^–1.Concentrations of most elements in leaf and twig were slightly greater in wetmonths than dry months, except for C and K. Woody litter had low N and Pconcentrations compared with the leaf and reproductive parts. Nutrient returntothe soil through small litterfall decrease in the orderC>N>Ca>K>Mg>Mn>Al>P>Fe, while nutrient reserve inlitteron the forest floor was in the declining sequenceC>N>Ca>K>Mg>P>Fe>Al>Mn.     

元江干热河谷植物叶片解剖和养分含量特征

研究了元江干热河谷旱田植物(旱季可浇灌,水分较好)和山坡半萨王纳植被中(自然状况,水分较差)共20种的叶片形态解剖特征,以及7种山坡植物叶片养分含量特征.结果表明,山坡植物叶片比叶重大,气孔密度大,气孔长度小,海绵组织/栅栏组织的值小等.元江干热河谷山坡植物叶片养分含量低,1.3%＞Ca＞N＞K＞1%＞Mg＞P＞S.除氮元素外,其它元素种间差别1～3倍.与热带植物群落叶片养元素含量相比,热带雨林＞元江山坡植物＞东南亚沙地旱生林和巴西矮卡廷加群落,表明元江干热河谷植物叶片具有明显的旱生性形态解剖特征,且叶片养分含量也较低.     

Effect of crop residues on root growth and phosphorus acquisition of pearl millet in an acid sandy soil in Niger

The effect of long-term (1983–1988) applications of crop residues (millet straw, 2–4 t ha^-1 yr^–1) and/or mineral fertilizer (30 kg N, 13 kg P and 25 kg K ha^-1 yr^-1) on uptake of phosphorus (P) and other nutrients, root growth and mycorrhizal colonization of pearl millet (Pennisetum glaucum L.) was examined for two seasons (1987 and 1988) on an acid sandy soil in Niger. Treatments of the long-term field experiment were: control (–CR–F), mineral fertilizer only (–CR+F), crop residues only (+CR–F), and crop residues plus mineral fertilizer (+CR+F).In both years, total P uptake was similar for +CR–F and –CR+F treatments (1.6–3.5 kg P ha^-1), although available soil P concentration (Bray I P) was considerably lower in +CR–F (3.2 mg P kg^-1 soil) than in –CR+F (7.4) soil. In the treatments with mineral fertilizers (–CR+F; +CR+F), crop residues increased available soil P concentrations (Bray I P) from 7.4 to 8.9 mg kg^-1 soil, while total P uptake increased from 3.6 to 10.6 kg P ha^-1. In 1987 (with 450 mm of rainfall), leaf P concentrations of 30-day-old millet plants were in the deficiency range, but highest in the +CR+F treatment. In 1988 (699 mm), leaf P concentrations were distinctly higher, and again highest in the +CR+F treatment. In the treatments without crop residues (–CR–F; –CR+F), potassium (K) concentrations in the leaves indicated K deficiency, while application of crop residues (+CR–F; +CR+F) substantially raised leaf K concentrations and total K uptake. Leaf concentrations of calcium (Ca) and magnesium (Mg) were hardly affected by the different treatments.In the topsoil (0–30 cm), root length density of millet plants was greater for +CR+F (6.5 cm cm^-3) than for +CR–F (4.5 cm cm^-3) and –CR+F (4.2 cm cm^-3) treatments. Below 30 cm soil depth, root length density of all treatments declined rapidly from about 0.6 cm cm^-3 (30–60 cm soil depth) to 0.2 cm cm^-3 (120–180 cm soil depth). During the period of high uptake rates of P (42–80 DAP), root colonization with vesicular-arbuscular mycorrhizal (VAM) fungi was low in 1987 (15–20%), but distinctly higher in 1988 (55–60%). Higher P uptake of +CR+F plants was related to a greater total root length in 0–30 cm and also to a higher P uptake rate per unit root length (P influx). Beneficial effects of crop residues on P uptake were primarily attributed to higher P mobility in the soil due to decreased concentrations of exchangeable Al, and enhancement of root growth. In contrast, the beneficial effect of crop residues on K uptake was caused by direct K supply with the millet straw.     

Lead dynamics in the forest floor and mineral soil in south-central Ontario

Recent studies have suggested that the residence time of Pb in the forest floor may not be as long as previously thought, and there is concern that the large pulse of atmospheric Pb deposited in the 1960s and early 1970s may move rapidly through mineral soils and eventually contaminate groundwater. In order to assess Pb mobility at a woodland (JMOEC) in south-central Ontario, a stable Pb isotope tracer ²⁰⁷Pb (8 mg m^–2) was added to the forest floor in white pine (Pinus strobus) and sugar maple (Acer saccharum) stands, respectively, and monitored over a 2-year period. Excess ²⁰⁷Pb was rapidly lost from the forest floor. Applying first-order rate coefficients (k) of 0.57 (maple) and 0.32 (pine) obtained from the tracer study, and estimates of Pb deposition in the region, current predicted Pb concentrations in the forest floor are 1.5–3.1 and 2.1–5.8 mg kg^–1 in the maple and pine plots, respectively. These values compare favorably with measured concentrations (corrected for mineral soil contamination) of 3.1–4.3 mg kg^–1 in the maple stand and 2.6–3.6 mg kg^–1 in the pine stand. The response time (1/^k) of Pb in the forest floor at the sugar maple and white pine plots was estimated to be 1.8 and 3.1 years, respectively. The rapid loss of Pb from the forest floor at the JMOEC is much greater than previously reported, and is probably due to the rapid rate of litter turnover that is characteristic of forests with mull-type forest floors. In a survey of 23 forested sites that border the Precambrian Shield in south-central Ontario, Pb concentrations in the forest floor increased exponentially with decreasing soil pH. Lead concentrations in the forest floor at the most acidic survey sites, which exhibited mor-type forest floors, were approximately 10 times higher (80 mg kg^–1) than at the JMOEC, and pollution Pb burdens were up to 25 times greater. Despite the rapid loss of Pb from the forest floor at the JMOEC, the highest pollution Pb concentrations were found in the upper (0–1 cm) mineral soil horizon. Lead concentrations in the upper 30 cm of mineral soil were strongly correlated with organic matter content, indicating that pollution Pb does not move as a pulse down the soil profile, but instead is linked with organic matter distribution, indicating groundwater contamination is unlikely.     

Responses of N fluxes and pools to elevated atmospheric CO2 in model forest ecosystems with acidic and calcareous soils

The objectives of this study were to estimate how soil type, elevated N deposition (0.7 vs. 7 g N m^–2y^–1) and tree species influence the potential effects of elevated CO₂ (370 vs. 570 mol CO₂ mol^–1) on N pools and fluxes in forest soils. Model spruce-beech forest ecosystems were established on a nutrient-rich calcareous sand and on a nutrient-poor acidic loam in large open-top chambers. In the fourth year of treatment, we measured N concentrations in the soil solution at different depths, estimated N accumulation by ion exchange resin (IER) bags, and quantified N export in drainage water, denitrification, and net N uptake by trees. Under elevated CO₂, concentrations of N in the soil solution were significantly reduced. In the nutrient-rich calcareous sand, CO₂ enrichment decreased N concentrations in the soil solution at all depths (–45 to –100%). In the nutrient-poor acidic loam, the negative CO₂ effect was restricted to the uppermost 5 cm of the soil. Increasing the N deposition stimulated the negative impact of CO₂ enrichment on soil solution N in the acidic loam at 5 cm depth from –20% at low N inputs to –70% at high N inputs. In the nutrient-rich calcareous sand, N additions did not influence the CO₂ effect on soil solution N. Accumulation of N by IER bags, which were installed under individual trees, was decreased at high CO₂ levels under spruce in both soil types. Under beech, this decrease occurred only in the calcareous sand. N accumulation by IER bags was negatively correlated with current-years foliage biomass, suggesting that the reduction of soil N availability indices was related to a CO₂-induced growth enhancement. However, the net N uptake by trees was not significantly increased by elevated CO₂. Thus, we suppose that the reduced N concentrations in the soil solution at elevated CO₂ concentrations were rather caused by an increased N immobilisation in the soil. Denitrification was not influenced by atmospheric CO₂ concentrations. CO₂ enrichment decreased nitrate leaching in drainage by 65%, which suggests that rising atmospheric CO₂ potentially increases the N retention capacity of forest ecosystems.     

Spatial and temporal patterns of root activity in a species-rich alluvial grassland

Summary The time and depth of activity of a number of co-existing grassland plants was measured using a technique involving the simultaneous injection to different depths in soil of 3 chemical tracers — Li, Rb and Sr. Root activity at a particular depth was assessed from the concentration of each tracer in leaf tissue.The seven most constant species showed very similar patterns of root activity, which was greater at 5 than at 15 or 25 cm except towards the end of the growth period in late June. Maximum root activity generally occurred earlier than maximum shoot productivity but there was little evidence of differentiation between species. When root activity was assessed as a proportion of total community root activity, by combining tracer concentration and biomass data, seasonal differences between species were more obvious. Using both root activity and productivity data, species were grouped into two main guilds, one active in spring (April-May) and one in summer (June).Correlations of above-ground biomass with root activity at different depths revealed that species of the spring guild were more active in the 5–15 cm horizons and those of the summer guild at 15–25 cm.These patterns suggest that rooting depth and time of activity are strongly linked: early-active species tend to be less productive and shallower-rooted and this combination of characters allows them to escape from competition with more productive species, by being active at a time when deeper soil layers are less hospitable.     

Leaf-consumption levels in subtropical mangroves of Paranaguá Bay (SE Brazil)

The objective of this study was to measure leaf consumption levels, mainly by insect herbivores and the tree-dwelling crab Aratus pisonii (H. Milne Edwards, 1837), in mangrove forests of a large subtropical estuarine system in the South Atlantic Ocean, to determine if patterns of herbivory varied with forest structure, tree species and marked seasonal differences in rainfall and temperature. We analyzed three structurally different mangroves located in the euhaline high-energy sector of Paranaguá Bay, all of them with known values of annual litter fall. Consumption levels varied from 2.2–5.4% of total leaf area considering each site as a whole, and from 2.0–6.0% considering each tree species separately. No significant differences in consumption levels were found between summer and winter samples, but significant differences were found among sites and among tree species. Leaves from Laguncularia racemosa were most consumed. The site with lower consumption levels was the one covered with dwarf trees, a condition usually caused by low nutrient availability in the soil. Analysis of nitrogen and phosphorus levels revealed lower amounts of both nutrients in soils and of phosphorus in leaves from this site when compared to the ones containing more developed trees. This result suggests a relationship between herbivory and nutrient availability in the plants.     

Leaf productivity along a precipitation gradient in lowland Panama: patterns from leaf to ecosystem

Moisture availability has the potential to affect tropical forest productivity at scales ranging from leaf to ecosystem. We compared data for leaf photosynthetic, chemical and structural traits of canopy trees, litterfall production and seasonal availability of soil water at four sites across a precipitation gradient (1,800–3,500 mm year^–1) in lowland Panamanian forest to determine how productivity at leaf and ecosystem scales may be related. We found stronger seasonality in soil water potential at drier sites. Values were close to zero at all sites during the wet season and varied between a minimum of –2.5 MPa and –0.3 MPa at the driest and wettest sites, respectively, during the dry season. Leaf photosynthesis and nitrogen concentration decreased with increasing precipitation, whereas leaf thickness increased with increasing precipitation. Leaf toughness and fiber/N ratios increased with increasing precipitation indicating reduced nutritional content and palatability with precipitation. Seasonality of litter production and quality decreased with increasing precipitation, but the amount of litterfall produced was not substantially different among sites. It appears that in Neotropical forest, moisture availability is associated with leaf photosynthetic and defensive traits that influence litterfall timing and quality. Therefore, variation in leaf physiological traits has the potential to influence decomposition and nutrient cycling through effects on litter quality.     

Litterfall and nutrient returns in red alder stands in western Washington

Summary Litterfall was collected over 1 year from eight natural stands of red alder growing on different sites in western Washington. The stands occurred at various elevations and on different soils, and differed in age, basal area, and site index. Most litterfall was leaf litter (average 86 percent). Amounts of litterfall and leaf litter varied significantly (P<0.05) among the sites. Average weights of litterfall and leaf litter in kg ha^–1 yr^–1, were 5150 and 4440, respectively. Weight of leaf litter was not significantly (P<0.05) related to site index, stand age, or basal area. The sites varied significantly (P<0.05) in concentrations of all elements determined in the leaf litter, except Zn. Average chemical concentrations were: N, 1.98 percent; P, 0.09 percent; K, 0.44 percent; Ca, 1.01 percent; Mg, 0.21 percent; S, 0.17 percent; SO₄–S, nil; Fe, 324 ppm; Mn, 311 ppm; Zn, 53 ppm; Cu, 13 ppm; and Al, 281 ppm. There were significant correlations between some stand characteristics and concentrations of some elements, and among the different chemical components of the leaf litter. Important correlations were found between stand age and P concentration (r=–0.84,P<0.01); weight of leaf litter and P concentration (r=0.74,P<0.05); weight of leaf litter and K concentration (r=0.71,P<0.05); concentrations of N and S (r=0.81,P<0.05); and concentrations of Fe and Al (r=0.98,P<0.01). Returns of the different elements to the soil by leaf litter varied among the different sites. Average nutrient and Al returns, in kg ha^–1 yr^–1, were: N, 82; Ca, 41; K, 19; Mg, 8; S, 7; P, 4; Fe, 1; Mn, 1; Al, 1; Zn, 0.2, and Cu, <0.1.     

Relationships between soil microbial properties and aboveground stand characteristics of conifer forests in Oregon

Eight forest sites representing a large range of climate, vegetation, and productivity were sampled in a transect across Oregon to study the relationships between aboveground stand characteristics and soil microbial properties. These sites had a range in leaf area index of 0.6 to 16 m² m^–2 and net primary productivity of 0.3 to 14 Mg ha^–1 yr^–1.Measurements of soil and forest floor inorganic N concentrations and in situ net N mineralization, nitrification, denitrification, and soil respiration were made monthly for one year. Microbial biomass C and anaerobic N mineralization, an index of N availability, were also measured. Annual mean concentrations of NH ₄ ⁺ ranged from 37 to 96 mg N kg^–1 in the forest floor and from 1.7 to 10.7 mg N kg^–1 in the mineral soil. Concentrations of NO ₃ ^– were low ( < 1 mg N kg^–1) at all sites. Net N mineralization and nitrification, as measured by the buried bag technique, were low on most sites and denitrification was not detected at any site. Available N varied from 17 to 101 mg N kg^–1, microbial biomass C ranged from 190 to 1230 mg Ckg^–1, and soil respiration rates varied from 1.3 to 49 mg C kg^–1 day^–1 across these sites. Seasonal peaks in NH ₄ ⁺ concentrations and soil respiration rates were usually observed in the spring and fall.The soils data were positively correlated with several aboveground variables, including leaf area index and net primary productivity, and the near infrared-to-red reflectance ratio obtained from the airborne simulator of the Thematic Mapper satellite. The data suggest that close relationships between aboveground productivity and soil microbial processes exist in forests approaching semi-equilibrium conditions.Abbreviations IR infrared - LAI leaf area index - k _c proportion of microbial biomass C mineralized to CO₂ - NPP net primary productivity - TM Thematic Mapper     

Environmental control of canopy dynamics and photosynthetic rate in the evergreen tussock grass Stipa tenacissima

Seasonal changes in leaf demography and gas exchange physiology in the tall evergreen tussock grass Stipa tenacissima, one of the few dominant plant species in the driest vegetation of Europe, were monitored over a period of two years at a field site in semi-arid south-eastern Spain. Three age-classes of leaves – young, mature and senescent – were distinguished in the green canopy. Production of new leaves and extension growth of older leaves occurred exclusively from October–November to May–June. The rate of extension was significantly correlated with gravimetric soil water content. Leaf growth ceased after gravimetric soil water content fell below 0.015 g g^–1 at the beginning of the dry season which corresponded to pre-dawn leaf water potentials of -3.0 MPa. Leaf senescence and desiccation reduced green leaf area by 43–49% during the dry season. Diurnal changes in the net photosynthetic rate of all three cohorts of leaves were bimodal with an early morning maximum, a pronounced midday depression and a small recovery late in the afternoon. Maximum photosynthetic rates of 10–16 mol CO₂ m^–2 s^–1 were attained from November 1993 to early May 1994 in young and mature leaves. Photosynthetic rate declined strongly during the dry season and was at or below compensation in September 1994. Gas exchange variables of young and mature leaves were not significantly different, but photosynthetic rate and diffusive conductance to water vapour of senescing leaves were significantly lower than in the two younger cohorts. Leaf nitrogen content of mature leaves varied seasonally between 2.9 and 5.2 g m^–2 (based on projected area of folded leaves), but was poorly correlated with maxima of the photosynthetic rate. There was a stronger linear relationship between the daily maxima of leaf conductance and pre-dawn leaf water potential than with atmospheric water vapour saturation deficit. Seasonal and between-year variation in daily carbon assimilation were caused mainly by differences in climatic conditions and canopy size whereas the effect of age structure of canopies was negligible. Since water is the most important limiting factor for growth and reproduction of S. tenacissima, any future rise in mean temperature, which might increase evapotranspiration, or decrease in rainfall, may considerably reduce the productivity of the grasslands, particularly at the drier end of their geographical distribution.     

三种苦苣苔对石灰土和红壤的适应性分析

为探讨苦苣苔科植物对其岩溶生境的适应性,该研究选取黄花牛耳朵(Primulina lutea)、紫花报春苣苔(Pri.purpurea)和桂林蛛毛苣苔(Paraboea guilinensis)三种苦苣苔科植物,将其栽种在石灰土及红壤两种不同类型的土壤中,观测记录其生长性状并对其叶片元素含量进行测定和比较。植株采集过程中,同时采集自然生境中三种苦苣苔科植物叶片及取样植物基部土壤,并对叶片及土壤元素的含量进行测定,作为今后苗圃试验的参照。结果表明:三种苦苣苔科植物在两种土壤上的生长状况及适应性具有差异,其在石灰土上生长良好,在红壤上生长较差;在两种不同土壤中,除N外,桂林蛛毛苣苔的叶片其他元素(P、K、Mn、Mg、Ca、Zn、Cu)差异极显著(P0.01);除P外,紫花报春苣苔的叶片其他元素(N、K、Mn、Mg、Ca、Zn、Cu)差异极显著(P0.01);除N、Cu、Ca外,黄花牛耳朵的叶片元素(P、K、Mn、Mg、Zn)差异极显著(P0.01);三种植物的叶片元素比值,除少数值没有差异外,大部分指标差异都极显著;对叶片元素与栽培土壤元素的相关性分析,发现植物叶片Mn元素与土壤中N、Ca、Mg、Zn、Mn、有机质含量等呈正相关,土壤P元素与叶片中N、P元素呈正相关,而与叶片中Zn元素呈负相关关系。在其他栽培条件一致的条件下,土壤因素及物种差别是造成黄花牛耳朵、紫花报春苣苔和桂林蛛毛苣苔适应性产生差异的主要原因。