Effects of topography on tropical lower montane forests under different geological conditions on Mount Kinabalu,Borneo

Species composition and forest structure change with topography.However, mechanisms for topographical vegetation changes are still not wellunderstood, because a topographical gradient is a complex environmentalgradientinclusive of many factors. The foot of Mt. Kinabalu is covered with three typesof geological substrates, i.e. Quaternary and Tertiary sedimentary rocks andultrabasic (serpentine) rock. Quaternary and Tertiary sedimentaryrocks are different in site age, but controlled in primary minerals. Tertiarysedimentary and ultrabasic rocks are contrasting in primary minerals, but arecomparable in age. This setting provides an opportunity to examine thevegetation differentiation along topographical gradients that are contrastinginmajor-nutrient supply due to the difference in site age and parent rock.We established a total of nine study plots by choosing three topographicalunits(ridge, middle- and lower-slope) on each substrate inthe tropical lower montane forest. Pool size and supply of soil N and Pdecreased upslope on each substrate, and the magnitude of the reduction fromslope to ridge decreased in the order of Quaternary sedimentary > Tertiarysedimentary > ultrabasic rock. Between-substrate difference in soilnutrient condition was greater on the lower-slopes than the ridges.Maximum tree size decreased and stem density increased upslope on eachsubstrate. Detrended correspondence analysis demonstrated that speciescomposition also changed along topographical gradients on all substrates.However, the magnitude of topographical changes in forest structure and speciescomposition varied with substrate and decreased from Quaternary sedimentary>Tertiary sedimentary > ultrabasic rock. The greatest between-substratedifference in vegetation occurred on the lower-slopes. Accordingly, ourresults suggest that the magnitude of vegetation changes due to topographybecomes smaller with decreasing pool size and supply of nutrients.     

Changes in biomass,productivity and decomposition along topographical gradients under different geological conditions in tropical lower montane forests on Mount Kinabalu,Borneo

We have examined how the structure and function of a forest ecosystem change with topography (lower-slope versus ridge) and how the changes are modified by nutrient availability depending on geological substrate (Quaternary and Tertiary sedimentary rocks and ultrabasic rock) in the tropical montane rain forests of Mt. Kinabalu (Borneo) where climate is humid and aseasonal. Reflecting the difference in site age and parent rock, the pool size of soluble-P and inorganic-N in topsoil decreased from Quaternary sedimentary >Tertiary sedimentary >ultrabasic rock on the lower-slope, and they decreased from the lower-slope to the ridge on all substrates. Forest structural attributes [stature, above-ground biomass, and leaf area index (LAI)] decreased in the order of Quaternary sedimentary >Tertiary sedimentary >ultrabasic rock in association with soil nutrients on the lower-slopes, and decreased upslope consistently on each of the three substrates. Functional attributes [above-ground net primary productivity (ANPP) and decomposition rate] demonstrated similar patterns to structure. ANPP significantly correlated with LAI among the six sites, while net assimilation rate (ANPP divided by LAI assuming an even productivity between above vs below-ground system) was nearly constant. Therefore, ANPP could be explained primarily by LAI. Topographical change in LAI could be explained by leaf mass per area (LMA) combined with stand-level leaf biomass. LMA increased upslope on all substrates in association with the decrease in individual leaf area. Stand-level leaf biomass decreased upslope on all substrates but the Tertiary sedimentary rock. Our study demonstrated that topography and geological substrates interactively affected forest structure and processes. The effect of topography on forest structure and processes was greater on nutrient-rich substrates than on poor substrates, and the effect of geological substrate was greater on lower-slopes than on ridges.     

Earthworm secondary production and N flux in agroecosystems: a comparison of two approaches

Production was estimated for Aporrectodea spp. and Lumbricus spp. populations in corn agroecosystems with a 5-year history of manure or inorganic fertilizer applications during 1994–1995 and 1995–1996. Earthworm biomass and production were greater in manure than inorganic fertilizer plots, although biomass and production declined by about 50% between 1994–1995 and 1995–1996 due to unfavorable climatic conditions. Production was highest during the spring and autumn when soil temperatures were between 4 and 22°C. Production was higher in Lumbricus spp. than Aporrectodea spp. populations due to greater Lumbricus spp. biomass. Aporrectodea spp. production was 3.47–16.14 g ash-free dry weight (AFDW) m^–2 year^–1, while Lumbricus spp. production was 6.09–18.11 g AFDW m^–2 year^–1, depending on the fertilizer treatment and the method used to estimate production. However, production estimates from the instantaneous growth rate method were within 27% of the values calculated using the size-frequency method. Nitrogen flux through earthworms was used to estimate efficiency quotients. Net production efficiency (P/A) ranged from 0.64 to 0.76, assimilation efficiency (A/C) ranged from 0.1 to 0.3, and gross production efficiency (P/C) ranged from 0.06 to 0.22. Annual N flux through earthworm populations was higher in manure than inorganic fertilizer plots, and ranged from 2.95 to 5.47 g N m^–2 year^–1 in 1994–1995 and 1.76 to 2.92 g N m^–2 year^–1 in 1995–1996. The N flux through earthworms represented an amount equivalent to 16–30% of crop N uptake during 1994–1995 and 11–18% of crop N uptake during 1995–1996. We concluded that the effects of earthworms on N cycling in corn agroecosystems were substantial, and that N flux through earthworms was influenced significantly by fertilizer amendments. Received: 20 September 1999 / Accepted: 24 March 2000     

Soil nitrogen mineralization rates of rainforests in a matrix of elevations and geological substrates on Mount Kinabalu, Borneo

Mt Kinabalu, Borneo, is characterized by a deep elevational gradient and mosaics of geological substrates. We chose a pair of two geological substrates (sedimentary vs ultrabasic) at five altitudes (800, 1400, 2100, 2700 and 3100 m a.s.l.). We investigated soil nitrogen (N) mineralization and nitrification rates using an incubation technique to assay the pattern and control of soil N status in this environmental matrix. In situ net mineralization rates decreased with elevation on both substrates. The decreasing pattern was linear across altitudes on ultrabasic rock, whereas on sedimentary rock it was depressed in the middle slope wet cloud zone. Sedimentary sites in this zone had low soil redox potential values and this anoxic soil condition might be related to slow N mineralization. The in situ rates were significantly greater (P < 0.05, anova) on sedimentary than on ultrabasic rock at the same altitudes except in the cloud zone. Net mineralization rates of the soils that were collected from different elevations and incubated in the same conditions were statistically invariable (P > 0.05) among the original elevations for sedimentary rock, but were variable (P < 0.05) for ultrabasic rock. Those of the soils that were collected from the same elevation and incubated at different elevations decreased significantly across altitudes (P < 0.05) for sedimentary rock, while they were invariable (P > 0.05) for ultrabasic rock. Thus, temperature had stronger effects on net N mineralization on sedimentary rock, whereas inherent soil quality had stronger effects on ultrabasic rock. Controls of soil N mineralization might be different between the two substrates, leading to diverse biogeochemical site conditions on Kinabalu.     

Gradational Forest Change along the Climatically Dry Valley Slopes of Bhutan in the Midst of Humid Eastern Himalaya

Altitudinal forest and climate changes from warm, dry valley bottom (1250 m a.s.l.) to cool, humid ridge top (3550 m a.s.l.) along the typical dry valley slopes of the Bhutan Himalaya were studied. Annual mean temperature decreased upslope with a lapse rate of 0.62 °C·100 m⁻¹ from 18.2 °C at the valley bottom to 4.3 °C at the ridge top. On the contrary volumetric soil moisture content increased from 14.7 to 75.0%. This inverse relationship is the major determinant factor for the distribution of different forest types along the altitudinal gradient. Based on the quantitative vegetation data from 15 plots arranged ca. 200 m in altitude interval (1520–3370 m a.s.l.), a total of 83 tree species belonging to 35 families were recorded. Three major formation types of lower and upper coniferous forests, and a mid-altitude evergreen and deciduous broad-leaved forest were contrasted. Including two transitional types, five forest zones were categorized based on cluster analysis, and each zone can be characterized by the dominants and their phytogeographical traits, viz. (1) west Himalayan warm, dry pine (1520–1760 m a.s.l.), (2) wide ranging east-west Himalayan mixed broad-leaved (1860–2540 m a.s.l.), (3) humid east Himalayan evergreen broad-leaved (2640–2820 m a.s.l.), (4) cool, humid east Himalayan conifer (2950–3210 m a.s.l.), and (5) wide ranging cold, humid conifer (3305–3370 m a.s.l.). Structurally, total basal area (biomass) increased from 15.2 m² ha⁻¹ in the pine forest (1520 m) to 101.7 m² ha⁻¹, in the conifer forest (3370 m a.s.l.). Similarly, soil organic carbon increased from 2.7 to 11.3% and nitrogen from 0.2 to 1.9% indicating dry, poor nutrient fragile ecosystem at the dry valley bottom. We concluded that low soil moisture content (<20%) limits downslope extension of broad-leaved species below 1650 m a.s.l. while coldest month’s mean temperature of −1 °C restricted the upslope extension of evergreen broad-leaved species above 3000 m a.s.l. Along the dry valley slopes, the transition from dry pine forest in the valley bottom, to a mixture of dry west Himalayan evergreen and deciduous east Himalayan broad-leaved, and to humid evergreen oak–laurel forests feature a unique pattern of forest type distribution.     

Soil-nutrient availability and the nutrient-use efficiencies of forests along an altitudinal gradient on Yakushima Island,Japan

The vegetation on Yakushima Island, Japan, grows on soils derived from Akahoya volcanic ash, released from the Kikai Caldera about 7300 years ago. The eruption was devastating and it is believed that primary succession and soil formation reinitiated across all altitudes at this point. We hypothesize that the concentrations of soil total phosphorus (P) and labile P fractions increase with increasing altitude because the soil formation has progressed less in upslope areas as a result of the cooler temperature and because of the ample P source of the volcanic ash. Conversely, we hypothesize that the concentration of soil inorganic nitrogen (N) decreases with increasing altitude. Available soil P and N would result in increasing P limitation downslope and increasing N limitation upslope, respectively. We studied soil P fractions and soil inorganic N, and P- and N-use efficiencies of the seven forests on Yakushima along an elevation gradient (170–1550 m a.s.l.). Contrary to our hypotheses, soil total P, labile soil P fractions, and inorganic N decreased with increasing altitude. The P- and N-use efficiencies of the forests were negatively correlated with the concentration of soil total active P (total P minus occluded P) and inorganic N, respectively. We suggest that progressive soil acidity and slower decomposition under cooler and wetter environments upslope must have dissolved the P contained in volcanic ash and accelerated P leaching. Forest ecosystems on Yakushima that show a distinct altitudinal zonation are, therefore, characterized by increasing P and N shortage with increasing altitude.     

The effect of calcium on flavanol production in cell suspension cultures of Polygonum hydropiper

Cultured Polygonum hydropiper cells maintained in Murashige and Skoog (MS) medium supplemented with 10^–6 m 2,4-D, 10^–6 m kinetin, 0.1% casamino acids and 3% sucrose were transferred to medium containing a higher concentration of calcium chloride (15 mm). The content of flavanols in the cells on the 6th day was approximately twice that of the control culture (31.9–60.7 mg/g dry wt). However, the contents of other secondary metabolites such as chlorogenic acid and gallic acid were not changed. The levels of flavanols in the culture medium remained unchanged throughout the 21-day culture period. Of the the inorganic components supplemented to the culture medium , only elevated levels of calcium chloride induced an increase in flavanol contents of the cells. The results indicated that the elevated concentration of calcium in the culture medium played an important role in activating the accumulation of flavanols. Received: 4 June 1998 / Revision received: 30 October 1998 / Accepted: 29 November 1998     

Hypereutrophication events in the Ca’Pisani lagoons associated with intensive aquaculture

Parameters of ecosystem structure and functioning were analyzed in three hypereutrophic lagoons of Ca’Pisani during the season of 2001. Lagoons are situated at wetlands of the NW Adriatic in the vicinity of the Porto Viro, Po River delta. They are associated with intensive fish culture enterprise and accept its wastewater. In June, the lagoons were found overloaded with the biomass of nitrophylic algae. At the end of July, a bloom of potentially toxic dinoflagellate Alexandrium tamarense occurred. Soon, it was supplanted by the picocyanobacterial assemblage, which arrived into the lagoons from the coastal Adriatic via the Marine channel. Wet biomass of this new picocyanobacterial bloom arrived in September attained 30–60 g m⁻³. Decrease of white disk water transparency down to 30–40 cm resulted in a gross mortality of macrophytes accompanied by spreading of floating saprobic alga Enteromorpha. Phytoplankton was dominated in June to July by small mixotrophic phytoflagellates with a wet biomass of 200–1300 mg m⁻³. Number of bacterioplankton ranged between 4 and 7 × 10⁶ ml⁻¹ and its wet biomass between 1.4 and 2.1 g m⁻¹. Its maximum of 18 × 10⁶ ml⁻¹ was observed in late August, when the mortality of macrophytes had occurred. Zooplankton and zoobenthos were found depleted in the lagoons especially during the blooms. Diel fluctuations of dissolved oxygen in the lagoons in June to July reached 150–200% of saturation. Photosynthetic oxygen production ranged between 15 and 30 g O₂ m⁻² d⁻¹. Water column deoxygenation rate was 1–1.5 mg O₂ l⁻¹ h⁻¹. Total photosynthesis production reached 3–8 g C m⁻² d⁻¹ by the share of phytoplankton 5–15%. Hyper-accumulation of total phosphorus in the water column and of toxic labile sulfides in the bottom sediments was documented. Content of inorganic phosphorus in water remained unusually high even by its intensive uptake by microplankton. The PO₄P uptake rate measured with ³²P-label ranged during the bloom of picocyanobacteria between 10 and 50 nM l⁻¹ min⁻¹, and the residence time of PO₄P between 15 and 50 min. The data were generalized via the calculation of energy balance and the deduction of the energy flow scheme in the ecosystem. Their analysis demonstrates the invalidation of ecosystems in hypereutrophic lagoons due to their overload with organic matter, with nutrients and with labile sulfides. After having depleted their animal food web, they are unable to decompose local plus external organic loading. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

青藏高原植被降水利用效率的空间格局及其对降水和气温的响应

植被降水利用效率(precipitation use efficiency, PUE)是反映生态系统水、碳循环相互关系的重要指标。该文利用GLOPEM-CEVSA模型模拟了青藏高原2000-2008年植被净初级生产力(net primary production, NPP), 以97个野外草地样点实测地上净初级生产力(above-ground net primary productivity, ANPP)对模拟NPP进行验证, 模拟NPP与ANPP线性显著相关(R ² = 0.49, p < 0.001)。利用降水量空间插值数据, 分析了近9年青藏高原植被PUE的空间分布、主要植被类型的PUE及其与降水量之间的变化关系。结果表明: 2000-2008年青藏高原地区植被年平均PUE沿东南向西北递减, 降水量和气温对植被PUE有着重要的影响; PUE在不同植被类型间差异较大, 其中农田PUE最高, 高寒草甸PUE高于高寒草原。在不同降水区域植被PUE与降水量的关系不同, 降水量低于90 mm的区域, 植被PUE值最低((0.026 ± 0.190) g C·m ^-2·mm ^-1, 平均值±标准偏差)、波动最大(变异系数CV = 721%), 与降水量和气温不相关(p = 0.38)。降水量为90-300 mm的地区, 植被PUE较低((0.029 ± 0.074) g C·m ^-2·mm ^-1, 平均值±标准偏差)、波动较大(CV = 252%), 与降水量和气温显著相关(p < 0.001), 降水量和气温能够解释PUE空间变化的43.4%, 其中降水量的影响是气温的1.7倍。降水量为300-650 mm的区域占整个研究区的45%, 主要植被类型为高寒草原, 植被PUE较高((0.123 ± 0.191) g C·m ^-2·mm ^-1, 平均值±标准偏差), CV为155%; 植被PUE的空间变化与降水量和气温极显著相关(p < 0.001), 降水量和气温能够解释植被PUE空间变化的97.8%, 但以气温影响为主导, 其影响是降水量的1.5倍。降水量为650 mm的区域, 植被PUE达到最高(0.26 g C·m ^-2·mm ^-1)。降水量为650-845 mm的区域主要是西藏林芝地区, 植被以常绿针叶林为主, PUE最高((0.210 ± 0.246) g C·m ^-2·mm ^-1, 平均值±标准偏差)、波动最小(CV = 117%); 降水量和气温可解释植被PUE空间变化的93.1% (p < 0.001), 降水量的影响是气温的3.5倍, 但其影响为负。     

Variation in soil phosphorus, sulfur, and iron pools among south Florida wetlands

To determine relationships between soil nutrient status and known gradients in primary production, we collected and analyzed soils from 17 LTER sampling sites along two transects through south Florida wetland ecosystems. Through upstream freshwater marsh, a middle reach including the oligohaline marsh/mangrove ecotone, and downstream estuarine habitats, we observed systematic variation in soil bulk density, organic content, and pools of phosphorus (P), inorganic sulfur, and extractable iron. Consistent with observed differences in wetland productivity known to be limited by P availability, total P averaged ~200 μg g dw⁻¹ in soils from the eastern Taylor Slough/Panhandle and was on average three times higher in soils from the western Shark River Slough. Along both transects, the largest pool of phosphorus was the inorganic, carbonate-bound fraction, comprising 35–44% of total P. Greater than 90% of the total inorganic sulfur pool in these south Florida wetland soils was extracted as pyrite. Freshwater marsh sites typically were lower in pyrite sulfur (0.2–0.8 mg g dw⁻¹) relative to marsh/mangrove ecotone and downstream estuary sites (0.5–2.9 mg g dw⁻¹). Extractable iron in freshwater marsh soils was significantly higher from the Taylor Slough/Panhandle transect (3.2 mg g dw⁻¹) relative to the western Shark River Slough transect (1.1 mg g dw⁻¹), suggesting spatial variation in sources and/or depositional environments for iron. Further, these soil characteristics represent the collective, integrated signal of ecosystem structure, so any long-term changes in factors like water flow or water quality may be reflected in changes in bulk soil properties. Since the objective of current Everglades restoration initiatives is the enhancement and re-distribution of freshwater flows through the south Florida landscape, the antecedent soil conditions reported here provide a baseline against which future, post-restoration measurements can be compared.     

Groundwater-induced redox-gradients control soil properties and phosphorus availability across four headwater wetlands, New York, USA

Hydrochemical patterns across groundwater-fed wetlands, especially carbonate and redox gradients, can influence phosphorus (P) availability by controlling its distribution among different soil pools. We explored these linkages by comparing shallow (5–20 cm) soil properties along groundwater flowpaths in two rich fens, a marl fen, and a poor fen. Organic matter content, bulk density, and total elemental content varied more with depth to underlying drift materials than with water table fluctuation, but also were influenced by groundwater discharge, which stabilized water table elevations and controlled redox conditions. Total sulfur and calcium content increased where pore-water chemistry indicated active iron and sulfate reduction. Calcium mineral dynamics, however, did not appear to influence P availability: first, calcium carbonate (CaCO₃) accounted for <2% of the soil composition, except in the marl fen where it accounted for 20–25% of the soil composition. Second, Ca-bound P pools, determined from hydrochloric extraction of wet soil samples, accounted for <25% of the inorganic soil P pool. In contrast, iron-bound P determined from bicarbonate-buffered dithionite solution, accounted for 50–80% of the inorganic soil P, and increased where there was evidence of groundwater mixing, as did P release rates inferred from incubated anion resin bags. The total carbon and phosphorus content of organic-rich soils as well as available and labile P pools were strongly correlated with pore-water iron and alkalinity concentrations. Groundwater discharge and resulting hydrochemical gradients explained significant variation in soil composition and P availability across each site. Results highlight the importance of conducting biogeochemical studies in the context of a site’s shallow geologic setting and suggest mechanisms supporting the diverse plant species unique to groundwater wetlands.     

Interactive influences of climate and parent material on soil microbial community structure in Bornean tropical forest ecosystems

Climate and parent material strongly control vegetation structure and function, yet their control over the belowground microbial community is poorly understood. We assessed variation in microbial lipid profiles in undisturbed forest soils (organic and surface mineral horizons) along an altitudinal gradient (700, 1,700, and 2,700 m a.s.l. mean annual temperature of 12–24°C) on two contrasting parent materials (acidic metasedimentary vs. ultrabasic igneous rock) in Mt. Kinabalu, Borneo. Soil organic carbon and nitrogen concentrations were generally higher at higher altitudes and, within a site, at upper soil horizons. Soil pH ranged from 3.9 to 5.3, with higher values for the ultrabasic soils especially at higher altitudes. The major shifts in microbial community structure observed were the decline in the ratio of fungal to bacterial lipid markers both with increasing soil depth and decreasing altitude. The positive correlation between this ratio with soil C and N concentrations suggested a strong substrate control in accord with the literature from mid to high-latitude ecosystems. Principal component analysis using seven groups of signature lipids suggested a significant altitude by parent material interaction—the significant difference in microbial community structure between the two rock types found at 2,700-m sites developed on weakly weathered soils diminished with decreasing altitude towards 700-m sites where soils were strongly weathered. These results are consistent with the hypothesis that parent material effect on soil microbial community (either directly via soil geochemistry or indirectly via floristic composition) is stronger at an earlier stage of ecosystem development.     

Stable soil nitrogen accumulation and flexible organic matter stoichiometry during primary floodplain succession

Large increases in nitrogen (N) inputs to terrestrial ecosystems typically have small effects on immediate N outputs because most N is sequestered in soil organic matter. We hypothesized that soil organic N storage and the asynchrony between N inputs and outputs result from rapid accumulation of N in stable soil organic pools. We used a successional sequence on floodplains of the Tanana River near Fairbanks, Alaska to assess rates of stable N accumulation in soils ranging from 1 to 500+ years old. One-year laboratory incubations with repeated leaching separated total soil N into labile (defined as inorganic N leached) and stable (defined as total minus labile N) pools. Stable N pools increased faster (2 g N m^–2 yr^–1) than labile N (0.4 g N m^–2 yr^–1) pools during the first 50 years of primary succession; labile N then plateaued while stable and total N continued to increase. Soil C pools showed similar trends, and stable N was correlated with stable C (r² = 0.95). From 84 to 95 % of soil N was stable during our incubations. Over successional time, the labile N pool declined as a proportion of total N, but remained large on an aerial basis (up to 38 g N m^–2). The stoichiometry of stable soil N changed over successional time; C:N ratios increased from 10 to 22 over 275 years (r² = 0.69). A laboratory ¹⁵N addition experiment showed that soils had the capacity to retain much more N than accumulated naturally during succession. Our results suggest that most soil N is retained in a stable organic pool that can accumulate rapidly but is not readily accessible to microbial mineralization. Because stable soil organic matter and total ecosystem organic matter have flexible stoichiometry, net ecosystem production may be a poor predictor of N retention on annual time scales.     

Production ecology of phyto- and zooplankton in a eutrophic pond dominated byChaoborus flavicans (Diptera: Chaobolidae)

Primary production of phytoplankton and secondary production of a daphnid and a chaoborid were studied in a small eutrophic pond. The gross primary production of phytoplankton was 290 gC m⁻² per 9 months during April–December. Regression analysis showed that the gross primary production was related to the incident solar radiation and the chlorophylla concentration and not to either total phosphorus or total inorganic nitrogen concentration. The mean chlorophylla concentration (14.2 mg m⁻³), however, was about half the expected value upon phosphorus loading of this pond. The mean zooplankton biomass was 1.60 g dry weight m⁻², of whichDaphnia rosea and cyclopoid copepods amounted to 0.69 g dry weight m⁻² and 0.61 g dry weight m⁻², respectively. The production ofD. rosea was high during May–July and October and the level for the whole 9 months was 22.6 g dry weight m⁻².Chaoborus flavicans produced 10 complete and one incomplete cohorts per year. Two consecutive cohorts overlapped during the growing season. The maximum density, the mean biomass, and the production were 19,100 m⁻², 0.81 g dry weight m⁻², and 11.7 g dry weight m⁻²yr⁻¹, respectively. As no fish was present in this pond, the emerging biomass amounted to 69% of larval production. The production ofC. flavicans larvae was high in comparison with zooplankton production during August–September, when the larvae possibly fed not only on zooplankton but also algae.     

Effects of labile soil carbon on nutrient partitioning between an arctic graminoid and microbes

We measured partitioning of N and P uptake between soil microorganisms and potted Festuca vivipara in soil from a subarctic heath in response to factorial addition of three levels of labile carbon (glucose) combined with two levels of inorganic N and P. The glucose was added to either non-sterilized or sterilized (autoclaved) soils in quantities which were within the range of reported, naturally occurring amounts of C released periodically from the plant canopy. The aims were, firstly, to examine whether the glucose stimulated microbial nutrient uptake to the extent of reducing plant nutrient uptake. This is expected in nutrient-deficient soils if microbes and plants compete for the same nutrients. Secondly, we wanted to test our earlier␣interpretation that growth reduction observed in graminoids after addition of leaf extracts could be caused directly by labile carbon addition, rather than by phytotoxins in the extracts. Addition of high amounts of N did not affect the microbial N pool, whereas high amounts of added P significantly increased the microbial P pool, indicating a luxury P uptake in the microbes. Both plant N and in particular P uptake increased strongly in response to soil sterilization and to addition of extra N or P. The increased␣uptake led to enhanced plant growth when both elements were applied in high amounts, but only led to increased tissue concentrations without growth responses when the nutrients were added separately. Glucose had strong and contrasting effects on plant and microbial N and P uptake. Microbial N and P uptake increased, soil inorganic N and P concentrations were reduced and plant N and P uptake declined when glucose was added. The responses were dose-dependent within the range of 0–450 μg C g⁻¹ soil added to the non-sterilized soil. The opposite responses of plants and microbes showed that plant acquisition of limiting nutrients is dependent on release of nutrients from the soil microbes, which is under strong regulation by the availability and microbial uptake of labile C. Hence, we conclude, firstly, that the microbial populations can compete efficiently with plants for nutrients to an extent of affecting plant growth when the microbial access to labile carbon is high in nutrient deficient soils. We also conclude that reduced growth of plants after addition of leaf extracts to soil can be caused by carbon-induced shifts in nutrient partitioning between plants and microbes, and not necessarily by phytotoxins added with the extracts as suggested by some experiments. Received: 15 February 1997 / Accepted: 12 July 1997     

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 NaHCO₃ 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.     

Treatment of swine manure effluent using freshwater algae: Production, nutrient recovery, and elemental composition of algal biomass at four effluent loading rates

Cultivating algae on nitrogen (N) and phosphorus (P) in animal manure effluents presents an alternative to the current practice of land application. The objective of this study was to determine how algal productivity, nutrient removal efficiency, and elemental composition of turf algae change in response to different loading rates of raw swine manure effluent. Algal biomass was harvested weekly from laboratory scale algal turf scrubber units using four manure effluent loading rates (0.24, 0.40, 0.62 and 1.2 L m⁻² d⁻¹) corresponding to daily loading rates of 0.3–1.4 g total N and 0.08–0.42 g total P. Mean algal productivity values increased from 7.1 g DW m⁻² d⁻¹ at the lowest loading rate (0.24 L m⁻² d⁻¹) to 9.4 g DW m⁻² d⁻¹ at the second loading rate (0.40 L m⁻² d⁻¹). At these loading rates, algal N and P accounted for> 90% of input N and 68–76% of input P, respectively. However, at higher loading rates algal productivity did not increase and was unstable at the highest loading rate. Mean N and P contents in the dried biomass increased 1.5 to 2.0-fold with increasing loading rate up to maximums of 5.7% N and 1.8% P at 1.2 L m⁻² d⁻¹. Biomass concentrations of Al, Ca, Cd, Fe, K, Mg, Mn, Mo, Si, and Zn increased 1.2 to 2.6-fold over the 5-fold range of loading rate. Biomass concentrations of Cd, K, Pb, and Si did not increase significantly with loading rate. At the loading rate of 0.40 L m⁻² d⁻¹ (corresponding to peak productivity) the mean concentrations of individual components in the algal biomass were (in mg kg⁻¹): 250 (Al), 4900 (Ca), 0.30 (Cd), 1050 (Fe), 3.4 (Pb), 2500 (Mg), 105 (Mn), 6.0 (Mo), 7,500 (K), and 510 (Zn). At these concentrations, heavy metals in the algal biomass would not be expected to reduce its value as a soil or feed amendment.     

Modelling the influence of bioturbation on the vertical distribution of sedimentary phosphorus in L. Esrom

A multilayer sediment-water exchange model was used to evaluate the importance of bioturbation in the profundal sediments of L. Esrom. The temporal variation of the vertical distribution of sedimentary phosphorus fractions was modelled with an objective function of 1.50. Deviations between measured and simulated values occurred in the spring, where the measured pool of sedimentary phosphorus sharply declined in the surface sediments. The application of a model for the activity ofChironomus anthracinus based on biomass, oxygen consumption and temperature improved the model in the spring period. The downwards transport of easy-degradable surface sediments reduced the average release of sedimentary phosphorus from 12 mg P · m⁻² · day⁻¹ to 11 mg P · M⁻² · day⁻¹. The introduction of a similar model for the other important burrowing species in L. Esrom,Potamothrix hammoniensis, lowered the objective function to 1.37 and increased the average release to 12.5 mg P · m⁻² · day⁻¹. The minor role of bioturbation in sediment processes is discussed.     

Soil phosphorus movement and budget after 13 years of fertilized cultivation in the Amazon basin

Agronomic studies on soil phosphorus dynamics have primarily focused on the plant-available inorganic P pool. However organic P and less labile inorganic pools can contribute significantly to plant P uptake. The objectives of this study were to determine the changes in inorganic and organic P pools of varying lability in and below the plowlayer after 13 years of continuous cultivation and fertilization on a Typic Paleudult in Yurimaguas, Peru. The field experiment was established after slash and burn of a secondary forest and included non-fertilized and fertilized treatments. The yearly cropping pattern consisted of an upland rice (Oryza sativa),-corn (Zea mays),-soybean (Glycine max) rotation. A modified version of the Hedley et al. procedure was used to sequentially fractionate soil P into increasingly recalcitrant organic and inorganic pools. Plowlayer accumulation of the fertilizer P occurred in all P pools. The greatest increase was in the NaOH extractable inorganic P pool. In the non-fertilized plots, the organic P decreased by 42%. Phosphorus fertilization resulted in significant movement of P below plowlayer. The accumulation occurred mostly in inorganic and organic P pools that are not quantified by traditional soil-P test methods. In fertilized plots sub-plowlayer total P increased by 90 g g^–1 (87%) while resin extractable P increased only 4 g g^–1. Phosphorus content of the organic P pools below the plowlayer increased by 24 g g^–1 (50%) in fertilized plots. The inclusion of less labile P pools in studies of P movement and the evaluation of P fertilizer residual values could lead to a better understanding of P dynamics and hence better management of P fertilization.     

Spatial and seasonal patterns of periphyton biomass and productivity in the northern Everglades,Florida, U.S.A.

We sampled periphyton in dominant habitats at oligotrophic and eutrophic sites in the northern Everglades during the wet and the dryseasons to determine the effects of nutrient enrichment on periphytonbiomass, taxonomic composition, productivity, and phosphorus storage. Arealbiomass was high (100–1600 g ash-free dry mass [AFDM]m⁻²) in oligotrophic sloughs and in stands of the emergentmacrophyte Eleocharis cellulosa, but was low in adjacent stands of sawgrass,Cladium jamaicense (7–52 g AFDM m⁻²). Epipelon biomasswas high throughout the year at oligotrophic sites whereas epiphyton andmetaphyton biomass varied seasonally and peaked during the wet season.Periphyton biomass was low (3–68 g AFDM m⁻²) and limitedto epiphyton and metaphyton in open-water habitats at eutrophic sites andwas undetectable in cattail stands (Typha domingensis) that covered morethan 90% of the marsh in these areas. Oligotrophic periphytonassemblages exhibited strong seasonal shifts in species composition and weredominated by cyanobacteria (e.g., Chroococcus turgidus, Scytonema hofmannii)during the wet season and diatoms (e.g. Amphora lineolata, Mastogloiasmithii) during the dry season. Eutrophic assemblages were dominated byCyanobacteria (e.g., Oscillatoria princeps) and green algae (e.g., Spirogyraspp.) and exhibited comparatively little seasonality. Biomass-specific grossprimary productivity (GPP) of periphyton assemblages in eutrophic openwaters was higher than for comparable slough assemblages, but areal GPP wassimilar in these eutrophic (0.9–9.1 g C m⁻²d⁻¹) and oligotrophic (1.75–11.49 g C m⁻²d⁻¹) habitats. On a habitat-weighted basis, areal periphytonGPP was 6- to 30-fold lower in eutrophic areas of the marsh due to extensiveTypha stands that were devoid of periphyton. Periphyton at eutrophic siteshad higher P content and uptake rates than the oligotrophic assemblage, butstored only 5% as much P because of the lower areal biomass.Eutrophication in the Everglades has resulted in a decrease in periphytonbiomass and its contribution to marsh primary productivity. These changesmay have important implications for efforts to manage this wetland in asustainable manner. This revised version was published online in July 2006 with corrections to the Cover Date.