Nelson T. Edwards

Separating root and soil microbial contributions to soil respiration: A review of methods and observations

Forest soil respiration is the sum of heterotrophic (microbes, soil fauna) and autotrophic (root) respiration. The contribution of each group needs to be understood to evaluate implications of environmental change on soil carbon cycling and sequestration. Three primary methods have been used to distinguish hetero- versus autotrophic soil respiration including: integration of components contributing to in situ forest soil CO2 efflux (i.e., litter, roots, soil), comparison of soils with and without root exclusion, and application of stable or radioactive isotope methods. Each approach has advantages and disadvantages, but isotope based methods provide quantitative answers with the least amount of disturbance to the soil and roots. Published data from all methods indicate that root/rhizosphere respiration can account for as little as 10 percent to greater than 90 percent of total in situ soil respiration depending on vegetation type and season of the year. Studies which have integrated percent root contribution to total soil respiration throughout an entire year or growing season show mean values of 45.8 and 60.4 percent for forest and nonforest vegetation, respectively. Such average annual values must be extrapolated with caution, however, because the root contribution to total soil respiration is commonly higher during the growing season and lower during the dormant periods of the year.

NET PRIMARY PRODUCTIVITY OF A CO2‐ENRICHED DECIDUOUS FOREST AND THE IMPLICATIONS FOR CARBON STORAGE

A central question concerning the response of terrestrial ecosystems to a changing atmosphere is whether increased uptake of carbon in response to increasing at- mospheric carbon dioxide concentration results in greater plant biomass and carbon storage or, alternatively, faster cycling of C through the ecosystem. Net primary productivity (NPP) of a closed-canopy Liquidambar styraciflua (sweetgum) forest stand was assessed for three years in a free-air CO2-enrichment (FACE) experiment. NPP increased 21% in stands ex- posed to elevated CO2, and there was no loss of response over time. Wood increment increased significantly during the first year of exposure, but subsequently most of the extra C was allocated to production of leaves and fine roots. These pools turn over more rapidly than wood, thereby reducing the potential of the forest stand to sequester additional C in response to atmospheric CO2 enrichment. Hence, while this experiment provides the first evidence that CO2 enrichment can increase productivity in a closed-canopy deciduous forest, the implications of this result must be tempered because the increase in productivity resulted in faster cycling of C through the system rather than increased C storage in wood. The fate of the additional C entering the soil system and the environmental interactions that influence allocation need further investigation.

Continuous Measurement of Carbon Dioxide Evolution From Partitioned Forest Floor Components

Loss rates of organic detritus in ecosystems can be determined by measuring rates of CO_2 release from decomposing organic substrates. We used an open system employing inverted boxes and an infrared gas analyzer (IRGA) to obtain continuous measurements of CO_2 evolution from the floor or a mixed mesophytic forest. Plexiglass sheets were used to partition respiratory activity of two litter horizons (01 and 02) and soil. Root respiration rates were determined with a differential respirometer. We compared the open system with a closed system in which KOH was used as the CO_2 absorber. Respiration measurements with KOH were 63% of IRGA values at 20 degrees C and 90% at 12 degrees. Flow rates (28 to 340 liters hr—1) had no effect on rates of CO_2 evolution. Nighttime CO_2 evolution rates were generally higher day—1. Preliminary data indicate annual CO_2 evolution of 3.8 kg m—2 (48% from litter, 17% from soil, and 35% from roots). This estimate is equivalent to 2.3 kg m—2 organic mass catabolized, assuming a carbohydrate substrate, which is 11% higher than independent estimates of the organic detritus annually available for catabolism.

Below-ground respiratory responses of sugar maple and red maple saplings to atmospheric CO2 enrichment and elevated air temperature

The research described in this paper represents a part of a much broader research project with the general objective of describing the effects of elevated [CO2] and temperature on tree growth, physiological processes, and ecosystem-level processes. The specific objective of this research was to examine the below-ground respiratory responses of sugar maple (Acer saccharum Marsh.) and red maple (Acer rubrum L.) seedlings to elevated atmospheric [CO2] and temperature. Red maple and sugar maple seedlings were planted in the ground in each of 12 open-top chambers and exposed from 1994 through 1997 to ambient air or air enriched with 30 Pa CO2,< in combination with ambient or elevated (+4 °C) air temperatures. Carbon dioxide efflux was measured around the base of the seedlings and from root-exclusion zones at intervals during 1995 and 1996 and early 1997. The CO2 efflux rates averaged 0.4 μmol CO2 m-2 s-1 in the root-exclusion zones and 0.75 μmol CO2 m-2 s-1 around the base of the seedlings. Mineral soil respiration in root-exclusion zones averaged 12% higher in the high temperature treatments than at ambient temperature, but was not affected by CO2 treatments. The fraction of total efflux attributable to root + rhizosphere respiration ranged from 14 to 61% in measurements made around red maple plants, and from 35 to 62% around sugar maple plants. Root respiration rates ranged from 0 to 0.94 μmol CO2 s-1 m-2 of soil surface in red maple and from 0 to 1.02 in sugar maple. In both 1995 and 1996 root respiration rates of red maple were highest in high-CO2 treatments and lowest in high temperature treatments. Specific red maple root respiration rates of excised roots from near the soil surface in 1996 were also highest under CO2 enrichment and lowest in high temperature treatments. In sugar maple the highest rates of CO2 efflux were from around the base of plants exposed to both high temperature and high-CO2, even though specific respiration rates were< lowest for this species under the high temperature and CO2 enrichment regime. In both species, patterns of response to treatments were similar in root respiration and root mass, indicating that the root respiration responses were due in part to differences in root mass. The results underscore the need for separating the processes occurring in the roots from those in the forest floor and mineral soil in order to increase our understanding of the effects of global climate change on carbon sequestration and cycling in the below-ground systems of forests.

Temperature-independent diet variations of respiration rates in Quercus alba and Liriodendron tulipifera

Diel cycles of bole and root respiration were observed in white oak and tulip poplar trees with highest rates occurring between 7 P.M. and midnight, and lowest rates between noon and 3 P.M. Calculated Q10 values, using daily mean temperatures and daily mean respiration rates, ranged from 1.9 to 4.8 and averaged 3.2 during several days in April and May. Respiration rates predicted from temperatures, in which Q10 was assumed equal to 2, followed a diel cycle 180 degrees out of phase with measured rates. It is suggested that any use of the classical Q1o relationship to predict respiration rates in a natural forest environment be considered suspect unless other controlling variables are considered. Reducing sugar concentrations in tulip poplar boles followed a similar diel cycle as respiration rates, suggesting a correlation between reducing sugar concentrations and respiration rates. In girdling experiments, respiration rates immediately above the incision of a girdled tulip poplar were up to five times higher than the control, while rates below the girdle dropped to about one-third of the control by the twelfth week after the tree was girdled. Diel respiration cycles were observed both above and below the girdle. We suggest that trees possess a diel scheme of food utilization whereby food catabolism is synchronized with growth processes at night when moisture availability is high.

Biological responses of two soybean cultivars exposed to enhanced UVB radiation

A UVB exposure and monitoring system has been established at the Oak Ridge National Laboratorys Global Climate Change Research Facility. The system consists of a power supply, and data acquisition and exposure equipment to accomplish controlled, elevated exposure of terrestrial plants to UVB. Plant biomass, selected compounds that absorb UV radiation, and DNA integrity/damage were measured for two soybean cultivars [Glycine max (L.) Merr.] Forrest and Essex exposed to elevated UVB (32% above ambient) in this system. The biomass of each major plant organ was observed to be less in soybean cultivar Forrest upon exposure to enhanced UVB with the greatest response in seed pods and stems. In contrast, soybean cultivar Essex showed no biomass response to elevated UVB. Enhanced UVB caused significant (P < 0.1) changes in concentrations of UV-absorbing compounds in both soybean cultivars. The Essex cultivar had an increase in UV-absorbing compounds, whereas a decline was observed for soybean Forrest. There was a decrease in the integrity of DNA, as measured by strand breaks, from both cultivars at 30 and 52 days to exposure. DNA pyrimidine dimers in isolated plant DNA were measured with Micrococcus luteus UV endonuclease. DNA from soybean Forrest exposed to UVB and sampled at 30 and 52 days of exposure had significantly greater (P<0.05) pyrimidine dimer concentration (dimer frequency ≈ 1 dimer per 28,000 DNA bases) than either cultivar exposed to UV treatment for 1 day or Essex at days 30–52 (dimer frequencies < /1 per 120,000 bases of DNA). Decrease in DNA integrity and biomass production in Forrest under elevated UVB may be related to the inability to maintain high concentrations of UV-absorbing compounds in leaves. The tolerant cultivar Essex increased the concentratio of UV-absorbing compounds while maintaining biomass production and DNA integrity under elevated UVB.

Growth, physiology, and nutrition of loblolly pine seedlings stressed by ozone and acidic precipitation : a summary of the ropis-south project

Previously published results from a multidisciplinary research program, Response of Plants to Interacting Stress (ROPIS), initiated by the Electric Power Research Insitute are summarized here. The overall objective of the ROPIS program was to develop a general mechanistic theory of plant response to air pollutants and other stresses. Direct and indirect phytotoxic impacts of O3 combined with induced deficiencies of key nutrients as a consequence of acidic deposition are important components in many of the hypotheses used to explain reported declines in forest growth. In order to address these concerns as they relate to loblolly pine (Pinus taeda L.) growth and develop a greater level of mechanistic understanding of stress response, a study was formulated with two major objectives: (i) over a multi-yr period evaluate the role of loblolly pine genotype in governing loblolly growth response to O3; and (ii) determine the underlying physiological and edaphic basis for loblolly growth response to O3, acidic precipitation, and soil Mg status. An open-top chamber facility located at Oak Ridge, TN provided controlled O3 exposure for the genotype screening study (1986–88) and controlled O3 exposure and rainfall exclusion and addition for the O3-rainfall acidity-soil Mg interaction study (1987–89). A variety of experimental techniques, measurements, and statistical procedures were used over a 4-yr period to quantify various aspects of plant growth, physiology, and soil-plant relationships. Results from the genotype screening study indicate that although family-specific O3 effects were observed at the end of the first year, no statistically significant O3 effects on diameter, height, or total biomass were evident at the end of three growing seasons; nor were any significant O3-family interactions found. In the interaction study, rainfall acidity and soil Mg level had only minimal affects on seedling growth and physiology. Ozone exposure produced significant changes in many variables, the most important being a net retention of carbon in above-ground biomass and a subsequent reduction in carbon allocation to the root system. This change could have important longterm implications for the trees ability to obtain water and nutrients, maintain important rhizosphere organisms, and achieve a level of vigor that protects against disease and insect attack.

Growth of Pinus Taeda L. Seedlings varies with Family and ozone exposure level

Loblolly pine seedlings of five half-sib families were grown under ambient, subambient (approximately 0.6 × ambient), and elevated [ambient + 60 ppb O3 (120 μg m−3)] O3 levels for one growing season in open-topped chambers. Diameter and height of the seedlings were measured periodically over the growing season, and above ground and below ground biomass were determined at harvest. Significant genetic differences were found in above ground volume (D2H) 1 mo after 03 fumigation began and continued until harvest. Biomass of secondary needles and coarse and fine roots also differed significantly among families. Elevated O3 resulted in significantly decreased D2H and secondary needle biomass relative to seedlings grown in ambient air. Seedlings receiving subambient O3 levels were intermediate in size, but were not significantly different from seedlings fumigated at ambient O3 levels. Root and stem biomass did not differ significantly among treatments. A significant interaction of O3 and genotype was detected, suggesting that the response of loblolly pine to O3 is influenced by genotype.

Field performance of the Walker Branch throughfall displacement experiment

The authors are conducting a large-scale manipulative field experiments in an upland oak forest on the Walker Branch Watershed in eastern Tennessee USA to identify important ecosystem responses that might result from future precipitation changes. The manipulation of soil moisture is being implemented by a gravity-driven transfer of throughfall precipitation from one treatment plot to another. Throughfall is intercepted in {approx} 2,000 subcanopy troughs (0.3 x 5 m) suspended above the forest floor of the dry plots ({approx} 33% of the ground area is covered) and transferred by gravity flow across an ambient plot for subsequent distribution onto the wet treatment plot. Percent soil water is being monitored with time domain reflectometers at 310 sampling locations across the site. The experimental system is able to produce statistically significant differences in soil water content in years having both extremely dry and extremely wet conditions. Furthermore, comparisons of pre- and post-installation soil temperature measurements have documented the ability of the experimental design to produce these changes without changing the microclimate of the forest understory.

Three-year growth responses of Pinus taeda L. to simulated rain chemistry, soil magnesium status, and ozone

Height, diameter, and biomass were measured for loblolly pine (Pinus taeda L) seedlings grown in soil containing 15 or 35 Μg Mg g−1 and exposed from May to October in 1987, 1988, and 1989 to three O3 concentrations (sub-ambient, ambient, or twice-ambient) and to rain pH levels of 3.8 or 5.2. Reduction in biomass accumulation in seedlings exposed to twice-ambient O3vs sub-ambient O3 was 14% (P = 0.03) in 1987, 11.4% (P = 0.002) by 1988, and 8% (P = 0.15) by 1989. The greatest height growth occurred in seedlings exposed to twice-ambient O3, and the greatest stem diameter growth occurred in seedlings exposed to sub-ambient O3. A comparison of stem volume (d2h) with stem biomass suggested that tissue density was reduced by elevated O3. Biomass accumulation response to rainfall chemistry was small (5.5% reduction in the low pH treatment in 1989) and not statistically significant for most plant tissues. Growth response to soil Mg status was not significant. Hoewever, in 1989 treatment interactions between rainfall chemistry and soil Mg status were observed. Height was 5% greater (P = 0.02) and biomass was 6% greater (P = 0.10) in seedlings grown in higher-Mg soil and receiving higher-pH rainfall than seedlings grown in any of the other pH-Mg treatment combinations. The data suggest direct adverse effects of near ambient O3 and indirect, slower acting and interacting adverse effects of rainfall chemistry and soil nutrient status on growth of loblolly pine.