Deuterium or tritium labeled water traditionally has been used for the measurement of total body water by application of the dilution principle. However, these methods have not enjoyed wide clinical use. The use of deuterium is hampered by the tedious and time consuming nature of the analysis while the use of tritium involves a radiation hazard. In addition, exchange of the label with nonaqueous hydrogen in the body raises questions about the accuracy of total body water values. To determine if water labeled with the stable isotope 18O can yield faster and more accurate results, total body water was measured simultaneously using water labeled with both 18O and 2H. The 18O and 3H dilutions were measured by mass spectrometry. The relative precision of the body water value using the 18O method was 2% for both serum and breath analysis. The 18O was fully equilibrated within 2 to 3 hr after administration, and results from the analysis of breath CO2 could be readily obtained within 1 hr after sampling. The H2(18)O dilution space averaged 3.0% (SE = 0.4) less than the 2HHO dilution space, because the latter exchanges with nonaqueous hydrogen. For this reason, the H2(18)O dilution should be a more accurate measure of total body water than the 3HHO dilution.
Although it is well known that fire can exert strong control on stand structure, composition, and dynamics in savannas and woodlands, the relationship between fire frequency and stand structure has been characterized in few of the world's savanna and woodland ecosystems. To address this issue in temperate oak-dominated ecosystems, we studied the effects of fire frequency on stand structure and dynamics in oak savanna and woodland stands that had been burned 0-26 times in 32 yr, in the Anoka Sand Plain region of Minnesota (USA).Seedling densities declined with increasing fire frequency, but differentially, for northern pin oak (Quercus ellipsoidalis), black cherry (Prunus serotina), serviceberry (Amelanchier sp.), and red maple (Acer rubrum). Bur oak (Q. macrocarpa) seedling density was not sensitive to fire frequency. Frequent burning (at least three fires per decade) prevented development of a sapling layer and canopy ingrowth. Low-frequency burning (fewer than two fires per decade) produced stands with dense sapling thickets.Reductions in overstory density and basal area from 1984 to 1995 were observed for all stands burned two or more times during that period. Basal area declined by 4-7% per year, and density declined by 6-8% per year in stands burned four or more times. Mortality rates in burned stands were higher for northern pin oak (50%) than for bur oak (8%). Northern pin oak mortality was highest for small trees (Ͻ 20 cm dbh) and lowest for mature trees (30-40 cm dbh); mortality increased with fire frequency. Bur oak mortality declined with increasing fire frequency.Attempts to preserve and maintain savannas as a viable ecosystem type in this region will require a long-term commitment to restoration-based management, with prescribed fire as a central tool. Burn frequency treatments with four or more fires per decade produce similar reductions in stem density and stand basal area but may lead to unsustainable oak tree populations. Within this general range, fire frequencies at a decadal scale should be chosen to address other management objectives, including suppressing shrubs and promoting increased cover of grasses and other herbaceous species. Fire management with a long-term view may also require periodic respites to allow for new cohorts of mature oak trees.
Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.
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