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Solar equivalent leaf area: an efficient biometrical parameter of individual leaves, trees and stands.

The solar equivalent leaf area (A(s)), a simply and easily determined biometrical parameter of leaves, trees and stands, was derived theoretically. The parameter is defined as projected leaf area weighted for the time integral of irradiance at a given location in the canopy relative to that of fully irradiated leaves at the top of the canopy. The efficiency of A(s) as a basis for estimating stand-area transpiration of a mature oak (Quercus robur L.) forest from measurements of transpiration by individual trees was compared with that of other stand and tree characteristics. Stand transpiration estimates based on A(s) were more precise and less prone to systematic error than estimates based on basal area, timber volume, projected tree crown area, projected leaf area, or leaf dry mass. Solar equivalent leaf area reflects both the amount and the physiological properties of leaves and can be used as a measure of tree size and functional capacity. It can be calculated from ordinary forest inventory data on trees and stands, adjusted according to simple phyllometric data. It appears to have wide application in ecological and forestry studies for relating the physiological characteristics of individual leaves to those of entire trees or stands.

Journal Article↗

Estimation of leaf area index in fourteen southern Wisconsin forest stands using a portable radiometer.

Projected leaf area index (LAI) and Beer-Lambert Law extinction coefficients (K) were estimated for 28-year-old Picea abies (L.) Karst., Larix decidua Mill., Pinus resinosa Ait., and Pinus strobus L. plantations using vertical profile data obtained with a portable integrating radiometer (sunfleck ceptometer). Predicted LAI values were compared with direct measures of LAI. Based on dimensional analysis, LAI ranged from 5.0 for Larix decidua to 10.5 for Picea abies. Significant inverse relationships between cumulative LAI and canopy transmitted radiation were observed for the four species (R(2) = 0.92-0.97). Beer-Lambert extinction coefficients ranged from 0.39 for Picea abies to 0.84 for Pinus strobus. Stand-level predictions of LAI based on the Beer-Lambert Law were compared with measured LAI values for eight conifer and six broadleaf stands. Using local K estimates resulted in predicted LAI values with an average 6% error. Using published K values resulted in an average error of 20%. High LAI and concomitantly low light levels below the canopy of Picea abies stands resulted in large overestimation errors in predicted LAI, rendering the sunfleck ceptometer inappropriate for forests with large LAIs.

Journal Article↗

Reconstructed development of stem production and foliage mass and its vertical distribution in Japanese larch.

Hypotheses of independence with time in (1) height growth rate, (2) crown length, (3) total foliage mass, and (4) vertical profile of foliage mass density within a crown following canopy closure were tested with two codominant trees in a 15-year-old Japanese larch (Larix kaempferi (Lamb.) Carr.) plantation by means of a combined method consisting of detailed stem analysis (to reconstruct development of stem growth over time), knot dissection (to identify time of branch mortality), and a stem diameter versus cumulative foliage mass relationship at a given stem location. I failed to reject all hypotheses at the 5% significance level. All parameters converged to constant values after canopy closure. Foliage efficiency for stem volume density production also assumed a constant value. Vertical profiles of the foliage mass density did not change after crown closure; however, the profile for one tree showed a more heterogeneous distribution because of the presence of large branches. Implications of these results for the profile diagram model of tree growth (Chiba et al. 1988, Osawa et al. 1990) are discussed.

Journal Article↗

Modeling changes in red spruce carbon balance and allocation in response to interacting ozone and nutrient stresses.

The simulation model TREGRO was developed to analyze the response of red spruce saplings to multiple stresses, such as drought, nutrient deficiency, and exposure to pollutants. The model provides a method of identifying changes in structural and non-structural carbon resources in the tree that may become measurable only after many years of exposure. The model is based on the assumption that the ability of plants to take up and use carbon, water, and nutrients depends on the interrelationships in availability among the three resources. Consequently, the model simulates the simultaneous cycling of these resources. In the model, the tree is divided into the following compartments: a canopy of leaves grouped by age class, branches, stem, and coarse and fine roots in a number of soil horizons. In each of these compartments we track three carbon pools: living structure, dead structure or wood, and total non-structural carbohydrate. The model calculates the photosynthesis of an entire red spruce tree each hour as a function of ambient environmental conditions and the availability of light, water, and nutrients; the daily redistribution of carbon throughout the plant; and the loss of carbon by respiration and senescence. To accomplish this task, the model tracks the flow of carbon dioxide to the sites of fixation within the leaves, the availability of light in the canopy, water and nutrient resources in each of three soil horizons, and the amounts of these resources taken up by the tree. Soil and plant water potentials, photosynthesis, and leaf respiration are simulated on an hourly timestep; nutrient uptake, allocation and growth are computed on a daily timestep. Through a set of example simulations, we demonstrate how the model can be used to examine the mechanisms by which plants respond to stresses experienced alone and in combination. The model was used to predict the growth decrease and the shifting pattern of carbon allocation expected for an isolated tree exposed to ozone and decreased nutrient availability due to acidic deposition. Decreased nutrient availability resulted in decreased growth and preferential carbon allocation to roots, which helped to alleviate the nutrient stress. Ozone stress also resulted in decreased plant growth but had the opposite effect on allocation patterns, with most of the growth reduction occurring in roots. The effect of simultaneous ozone and nutrient stress on tree growth was less than the sum of the independent single stresses, contrary to our expectation. This modeling approach can aid in evaluating the long-term effect of stress on resource availability, the potential for gradual deterioration of tree health under long periods of stress, and imbalances in growth accompanying shifts in carbon allocation caused by stress.

Journal Article↗

A simple method for estimating gross carbon budgets for vegetation in forest ecosystems.

Gross carbon budgets for vegetation in forest ecosystems are difficult to construct because of problems in scaling flux measurements made on small samples over short periods of time and in determining belowground carbon allocation. Recently, empirical relationships have been developed to estimate total belowground carbon allocation from litterfall, and maintenance respiration from tissue nitrogen content. I outline a method for estimating gross carbon budgets using these empirical relationships together with data readily available from ecosystem studies (aboveground wood and canopy production, aboveground wood and canopy biomass, litterfall, and tissue nitrogen contents). Estimates generated with this method are compared with annual carbon fixation estimates from the Forest-BGC model for a lodgepole pine (Pinus contorta Dougl.) and a Pacific silver fir (Abies amabilis Dougl.) chronosequence.

Journal Article↗

Influence of thermoregulatory vasomotion and ambient temperature variation on the accuracy of core-temperature estimates by cutaneous liquid-crystal thermometers.

BACKGROUND: Recently, liquid crystal skin-surface thermometers have become popular for intraoperative temperature monitoring. Three situations during which cutaneous liquid-crystal thermometry may poorly estimate core temperature were monitored: (1) anesthetic induction with consequent core-to-peripheral redistribution of body heat, (2) thermoregulatory vasomotion associated with sweating (precapillary dilation) and shivering (minimal capillary flow), and (3) ambient temperature variation over the clinical range from 18-26 degrees C. METHODS: The core-to-forehead and core-to-neck temperature difference was measured using liquid-crystal thermometers having an approximately 2 degrees C offset. Differences exceeding 0.5 degree C (a 1 degree C) temperature range) were a priori deemed potentially clinically important. Seven volunteers participated in each protocol. First, core-to-peripheral redistribution of body heat was produced by inducing propofol/desflurane anesthesia; anesthesia was then maintained for 1 h with desflurane. Second, vasodilation was produced by warming unanesthetized volunteers sufficiently to produce sweating; intense vasoconstriction was similarly produced by cooling the volunteers sufficiently to produce shivering. Third, a canopy was positioned to enclose the head, neck, and upper chest of unanesthetized volunteers. Air within the canopy was randomly set to 18, 20, 22, 24, and 26 degrees C. RESULTS: Redistribution of body heat accompanying induction of anesthesia had little effect on the core-to-forehead skin temperature difference. However, the core-to-neck skin temperature gradient decreased approximately 0.6 degree C in the hour after induction of anesthesia. Vasomotion associated with shivering and mild sweating altered the core-to-skin temperature difference only a few tenths of a degree centigrade. The absolute value of the core-to-forehead temperature difference exceeded 0.5 degree C during approximately 35% of the measurements, but the difference rarely exceeded 1 degree C. The core-to-neck temperature difference typically exceeded 0.5 degree C and frequently exceeded 1 degree C. Each 1 degree C increase in ambient temperature decreased the core-to-fore-head and core-to-neck skin temperature differences by less than 0.2 degree C. CONCLUSIONS: Forehead skin temperatures were better than neck skin temperature at estimating core temperature. Core-to-neck temperature differences frequently exceeded 1 degree C (a 2 degrees C range), whereas two thirds of the core-to-forehead differences were within 0.5 degree C. The core-to-skin temperature differences were, however, only slightly altered by inducing anesthesia, vasomotor action, and typical intraoperative changes in ambient temperature.

Adult↗

Measurement of gas exchange in intensive care: laboratory and clinical validation of a new device.

The performance of a new gas exchange monitor was assessed both in laboratory simulation and in ICU patients. Laboratory simulation using N2 and CO2 injections resulted in a mean error of 2 +/- 2% in CO2 production (VCO2) and 4 +/- 4% in oxygen consumption (VO2) in respirator measurements (n = 55) and in a mean error of 3 +/- 2% in VCO2 and 4 +/- 2% in VO2 in canopy measurements (n = 25). The mean error in RQ during ethanol burning was 2 +/- 2% in respirator measurements (n = 45) and 1 +/- 1% in canopy measurements. FIO2 had little effect on the accuracy of VCO2, whereas the accuracy on high rates of VO2 (VO2 = 400 ml/min) was reduced, when FIO2 increased: the error ranged from 1 +/- 1% to 6 +/- 1%, except at VO2 400 ml/min during FIO2 0.8, where the error was 16 +/- 3%. Neither peak airway pressure (+13 to +63 cm H2O) nor PEEP (0 to +20 cm H2O) had an effect on the accuracy. The highest level of minute ventilation studied (22.5 L/min) reduced the accuracy slightly (mean error of VCO2 4 +/- 1% and VO2 7 +/- 2%). In patients during controlled mechanical ventilation, increasing FIO2 from 0.4 to 0.6 had no effect on the results. VO2 was consistently higher by gas exchange than by the Fick principle: 16 +/- 9% during controlled ventilation (n = 20), 21 +/- 8% on synchronized intermittent mandatory ventilation (n = 10) and 25 +/- 8% during spontaneous breathing. We conclude that the device proved to be accurate for gas exchange measurements in the ICU.

Analysis of Variance↗

Falling palm fronds structure amazonian rainforest sapling communities.

The senescence and loss of photosynthetic and support structures is a nearly universal aspect of tree life history, and can be a major source of disturbance in forest understoreys, but the ability of falling canopy debris in determining the stature and composition of understorey communities seems not to have been documented. In this study, we show that senescent fronds of the palm Iriartea deltoidea cause substantial disturbance in tropical forest sapling communities. This disturbance influences the species composition of the canopy and subcanopy by acting as an ecological filter, favouring sapling species with characteristics conducive to recovery after physical damage. The scale of this dominance suggests that falling I. deltoidea debris may be influencing sapling community structure and species composition in Amazonian rainforests over very large spatial scales.

Analysis of Variance↗

Genetic rescue of remnant tropical trees by an alien pollinator.

Habitat fragmentation is thought to lower the viability of tropical trees by disrupting their mutualisms with native pollinators. However, in this study, Dinizia excelsa (Fabaceae), a canopy-emergent tree, was found to thrive in Amazonian pastures and forest fragments even in the absence of native pollinators. Canopy observations indicated that African honeybees (Apis mellifera scutellata) were the predominant floral visitors in fragmented habitats and replaced native insects in isolated pasture trees. Trees in habitat fragments produced, on average, over three times as many seeds as trees in continuous forest, and microsatellite assays of seed arrays showed that genetic diversity was maintained across habitats. A paternity analysis further revealed gene flow over as much as 3.2 km of pasture, the most distant pollination precisely recorded for any plant species. Usually considered only as dangerous exotics, African honeybees have become important pollinators in degraded tropical forests, and may alter the genetic structure of remnant populations through frequent long-distance gene flow.

Animals↗

Growth and survivorship of dipterocarp seedlings: differences in shade persistence create a special case of dispersal limitation.

A series of growth experiments and observations on natural populations have been carried out on dipterocarp species of contrasting ecology growing in artificial gaps and the forest understorey. These studies have demonstrated that although differences exist between species in photosynthetic and growth responses to the high-light environment, competition for light in canopy gaps is highly asymmetrical and tends to reinforce any pre-existing dominance hierarchy. We propose that differences in seedling persistence in forest canopy shade are highly influenced by species-specific biotic and abiotic interactions. Our experiments suggest that as seedlings, dipterocarp species trade off traits which enhance persistence and growth in shade against those that enhance their ability to exploit gaps. Less competitive species survive for progressively longer periods of time after a gregarious fruiting event. This leads to significant shifts with time in the number of species present in the seedling bank and hence in the importance of interspecific competition in determining which species dominates regrowth in gaps. We propose that this special case of dispersal limitation is more likely to account for coexistence of dipterocarp species than differences in growth responses to gaps of different size, with stochastic and environmental variables interacting to determine species distribution and abundance.

Ecosystem↗

Potential for ammonia recapture by farm woodlands: design and application of a new experimental facility.

There has been increasing pressure on farmers in Europe to reduce the emissions of ammonia from their land. Due to the current financial climate in which farmers have to operate, it is important to identify ammonia control measures that can be adopted with minimum cost. The planting of trees around farmland and buildings has been identified as a potentially effective and low-cost measure to enhance ammonia recapture at a farm level and reduce long-range atmospheric transport. This work assesses experimentally what fraction of ammonia farm woodlands could potentially remove from the atmosphere. We constructed an experimental facility in southern Scotland to simulate a woodland shelterbelt planted in proximity to a small poultry unit. By measuring horizontal and vertical ammonia concentration profiles within the woodland, and comparing this to the concentration of an inert tracer (SF6) we estimate the depletion of ammonia due to dry deposition to the woodland canopy. Together with measurements of mean ammonia concentrations and throughfall fluxes of nitrogen, this information is used to provide a first estimate of the fraction of emitted ammonia that is recaptured by the woodland canopy. Analysis of these data give a lower limit of recapture of emitted ammonia, at the experimental facility, of 3%. By careful design of shelterbelt woodlands this figure could be significantly higher.

Agriculture↗

Sampling ozone exposure of Canadian forests at different scales: some case studies.

The use of passive samplers in extensive monitoring, such as that used in national forest health monitoring plots, indicates that these devices are able to determine both spatial and temporal differences in ozone exposure of the plots. This allows for categorisation of the plots and the potential for cause-effect analysis of certain forest health responses. Forest exposure along a gradient of air pollution deposition demonstrates large variation in accumulated exposures. The efficacy of using passive samplers for in situ monitoring of forest canopy exposure was also demonstrated. The sampler data produced weak relationships with ozone values from the nearest "continuous" monitor, even though data from colocated samplers showed strong relationships. This spatial variation and the apparent effect of elevation on ozone exposure demonstrate the importance of topography and tree canopy characteristics in plant exposure on a regional scale. In addition, passive sampling may identify the effects of local pollutant gases, such as NO, which may scavenge ozone locally only to increase the production of this secondary pollutant downwind, as atmospheric reactions redress the equilibrium between concentrations of this precursor and those of the generated ozone. The use of passive samplers at the stand level is able to resolve vertical profiles within the stand and edge effects that are important in exposure of understorey and ground flora. Recent case studies using passive samplers to determine forest exposure to ozone indicate a great potential for the development of spatial models on a regional, landscape, and stand level scale.

Canada↗

Nitrogen deposition to and cycling in a deciduous forest.

The project described here seeks to answer questions regarding the role increased nitrogen (N) deposition is playing in enhanced carbon (C) sequestration in temperate mid-latitude forests, using detailed measurements from an AmeriFlux tower in southern Indiana (Morgan-Monroe State Forest, or MMSF). The measurements indicate an average atmosphere-surface N flux of approximately 6 mg-N m(-2) day(-1) during the 2000 growing season, with approximately 40% coming from dry deposition of ammonia (NH3), nitric acid (HNO3), and particle-bound N. Wet deposition and throughfall measurements indicate significant canopy uptake of N (particularly NH4+) at the site, leading to a net canopy exchange (NCE) of -6 kg-N ha(-1) for the growing season. These data are used in combination with data on the aboveground C:N ratio, litterfall flux, and soil net N mineralization rates to indicate the level of potential perturbation of C sequestration at this site.

Ammonia↗

Is there synchronicity in nitrogen input and output fluxes at the Noland Divide Watershed, a small N-saturated forested catchment in the Great Smoky Mountains National Park?

High-elevation red spruce [Picea rubens Sarg.]-Fraser fir [Abies fraseri (Pursh.) Poir] forests in the Southern Appalachians currently receive large nitrogen (N) inputs via atmospheric deposition (30 kg N ha(-1) year(-1)) but have limited N retention capacity due to a combination of stand age, heavy fir mortality caused by exotic insect infestations, and numerous gaps caused by windfalls and ice storms. This study examined the magnitude and timing of the N fluxes into, through, and out of a small, first-order catchment in the Great Smoky Mountains National Park. It also examined the role of climatic conditions in causing interannual variations in the N output signal. About half of the atmospheric N input was exported annually in the streamwater, primarily as nitrate (NO3-N). While most incoming ammonium (NH4-N) was retained in the canopy and the forest floor, the NO3-N fluxes were very dynamic in space as well as in time. There was a clear decoupling between NO3-N input and output fluxes. Atmospheric N input was greatest in the growing season while largest NO3-N losses typically occurred in the dormant season. Also, as water passed through the various catchment compartments, the NO3-N flux declined below the canopy, increased in the upper soil due to internal N mineralization and nitrification, and declined again deeper in the mineral soil due to plant uptake and microbial processing. Temperature control on N production and hydrologic control on NO3-N leaching during the growing season likely caused the observed inter-annual variation in fall peak NO3-N concentrations and N discharge rates in the stream.

Altitude↗

Atmospheric deposition of phosphorus to the everglades: concepts, constraints, and published deposition rates for ecosystem management.

This paper summarizes concepts underlying the atmospheric input of phosphorus (P) to ecosystems, published rates of P deposition, measurement methods, and approaches to future monitoring and research. P conveyed through the atmosphere can be a significant nutrient source for some freshwater and marine ecosystems. Particle sources and sinks at the land-air interface produce variation in P deposition from the atmosphere across temporal and spatial scales. Natural plant canopies can affect deposition rates by changing the physical environment and surface area for particle deposition. Land-use patterns can alter P deposition rates by changing particle concentrations in the atmosphere. The vast majority of P in dry atmospheric deposition is conveyed by coarse (2.5 to 10 m) and giant (10 to 100 m) particles, and yet these size fractions represent a challenge for long-term atmospheric monitoring in the absence of accepted methods for routine sampling. Most information on P deposition is from bulk precipitation collectors and wet/dry bucket sampling, both with questionable precision and accuracy. Most published annual rates of P deposition are gross estimates derived from bulk precipitation sampling in locations around the globe and range from about 5 to well over 100 mg P m(-2) year(-1), although most inland ecosystems receive between 20 and 80 mg P m(-2) year(-1). Rates below 30 mg P m(-2) year-1 are found in remote areas and near coastlines. Intermediate rates of 30 to 50 mg P m(-2) year(-1) are associated with forests or mixed land use, and rates of 50 to 100 mg P m(-2) year(-1) or more are often recorded from urban or agricultural settings. Comparison w ith other methods suggests that these bulk precipitation estimates provide crude boundaries around actual P deposition rates for various land uses. However, data screening cannot remove all positive bias caused by contamination of bucket or bulk collectors. As a consequence, continued sampling with these standard collectors in a region will not reduce the large uncertainty in rates derived from existing data. Calibrated surface accumulation methods hold promise as a primary means to estimate P flux in future monitoring. New methods for long-term P deposition monitoring will require an intercomparison of P flux estimates from surrogate surfaces, impactor sampling of particle concentrations combined with deposition models, and "throughfall" estimates for natural canopies. With better sampling methods and more long-term monitoring data, the importance of atmospheric P deposition in ecosystem dynamics and management can be better understood and predicted.

Atmosphere↗

A dynamic balance between gene activation and repression regulates the shade avoidance response in Arabidopsis.

Plants grown under dense canopies perceive through the phytochrome system a reduction in the ratio of red to far-red light as a warning of competition, and this triggers a series of morphological changes to avoid shade. Several phytochrome signaling intermediates acting as positive regulators of accelerated elongation growth and induction of flowering in shade avoidance have been identified. Here we report that a negative regulatory mechanism ensures that in the presence of far-red-rich light an exaggerated plant response does not occur. Strikingly, this unpredicted negative regulatory mechanism is centrally involved in the attenuation of virtually all plant responses to canopy shade.

Adaptation, Physiological↗

Concurrent comparisons of stomatal behavior, water status, and evaporation of maize in soil at high or low water potential.

Concurrent measurements of evaporation, leaf conductance, irradiance, leaf water potential, and osmotic potential of maize (Zea mays L. cv. Pa602A) in soil at either high or low soil water potential were compared at several hours on two consecutive days in July. Hourly evaporation, measured on two weighing lysimeters, was similar until 1000 hours Eastern Standard Time, but thereafter evaporation from the maize in the dry soil was always less than that in the wet soil; before noon it was 62% and by midafternoon, only 35% of that in the wet soil. The leaf water potential, measured with a pressure chamber, was between -1.2 and -2.5 bars and between -6.8 and -8 bars at sunrise (about 0530 hours Eastern Standard Time) in the plants in the wet and dry soil, respectively, but decreased quickly to between -8 and -13 bars in the plants in the wet soil and to less than -15 bars in the plants in the dry soil by 1100 to 1230 hours Eastern Standard Time. At this time, the leaf conductance of all leaves was less than 0.1 cm sec(-1) in the maize in the dry soil, whereas the conductance was 0.3 to 0.4 cm sec(-1) in the leaves near the top of the canopy in the wet soil. The osmotic potential, measured with a vapor pressure osmometer, also decreased during the morning but to a smaller degree than leaf water potential, so that by 1100 to 1230 hours Eastern Standard Time the leaf turgor potential was 1 to 2 bars in all plants. Thereafter, leaf turgor potential increased, particularly in the plants in soil at a high water potential, whereas leaf water potential continued to decrease even in the maize leaves with partly closed stomata. Evidently maize can have values of leaf conductance differing 3- to 4- fold at the same leaf turgor potential, which suggests that stomata do not respond primarily to bulk leaf turgor potential. Evidence for some osmotic adjustment in the plants at low soil water potential is presented. Although the degree of stomatal closure in the maize in dry soil did not prevent further development of stress, it did decrease evaporation in proportion to the decrease in canopy conductance.

Journal Article↗

Effects of Sink Removal on Photosynthesis and Senescence in Leaves of Soybean (Glycine max L.) Plants.

Photosynthetic rate, ribulose 1,5-bisphosphate carboxylase activity, specific leaf weight, and leaf concentrations of carbohydrates, proteins, chlorophyll, and inorganic phosphate were determined periodically from midbloom until maturity in leaves of soybean plants (Glycine max L., var. Hodgson) from which reproductive and vegetative sinks had been removed 32 hours before measurement, or continuously since midbloom.Leaf photosynthesis, measured in the top of the canopy, was partially inhibited by both sink removal treatments. This inhibition was of constant magnitude from midbloom until maturity.Leaf photosynthesis in the top of the canopy declined from midbloom until maturity in the control as well as in the desinked plants. The decline in photosynthesis was gradual at first, but later became more abrupt. The photosynthetic decline was equally evident in the yellowing leaves of control plants and in the dark green leaves of the continuously desinked plants.Neither the inhibition of photosynthesis by sink removal nor the decline in photosynthetic rate with time was clearly related to any of the measured traits.

Journal Article↗