Interactive effects of soil temperature and moisture on Concord grape root respiration

Root respiration has important implications for understanding plant growth as well as terrestrial carbon flux with a changing climate. Although soil temperature and soil moisture often interact, rarely have these interactions on root respiration been studied. This report is on the individual and combined effects of soil moisture and temperature on respiratory responses of single branch roots of 1-year-old Concord grape (Vitis labruscana Bailey) vines grown in a greenhouse. Under moist soil conditions, root respiration increased exponentially to short-term (1 h) increases in temperature between 10 degrees C and 33 degrees C. Negligible increases in root respiration occurred between 33 degrees C and 38 degrees C. By contrast to a slowly decreasing Q10 from short-term temperature increases, when roots were exposed to constant temperatures for 3 d, the respiratory Q10 between 10 degrees C and 30 degrees C diminished steeply with an increase in temperature. Above 30 degrees C, respiration declined with an increase in temperature. Membrane leakage was 89-98% higher and nitrogen concentration was about 18% lower for roots exposed to 35 degrees C for 3 d than for those exposed to 25 degrees C and 15 degrees C. There was a strong interaction of respiration with a combination of elevated temperature and soil drying. At low soil temperatures (10 degrees C), respiration was little influenced by soil drying, while at moderate to high temperatures (20 degrees C and 30 degrees C), respiration exhibited rapid declines with decreases in soil moisture. Roots exposed to drying soil also exhibited increased membrane leakage and reduced N. These findings of acclimation of root respiration are important to modelling respiration under different moisture and temperature regimes.     

Root respiration in citrus acclimates to temperature and slows during drought

Citrus seedlings were grown in soil columns in which the root system was hydraulically separated into two equal layers; this enabled us to maintain roots in the upper layer without water for 110 d. The columns were placed into waterbaths modified so that soil temperatures in the top layer could be maintained at 25°C or at 35°C, while temperature in the bottom layer was maintained at 25°C. We hypothesized that, if citrus plants were grown in dry soil for an extended period, root mortality would increase if the cost of maintaining the roots was increased by elevating the soil temperature. However, during the drought period we did not observe any root mortality, even at the higher soil temperature. Moreover, we did not find that root respiration was increased by prolonged exposure to drought and higher soil temperature. We did find that root respiration rates slowed in dry soil. Furthermore, when the soil columns were switched from one temperature treatment to another, root respiration rates in wet soil rapidly increased when moved to a higher temperature or rapidly decreased when moved to a lower temperature. But after only 4 d, respiration rates returned to their original level; root respiration in dry soil was not affected by either short-or long-term shifts in soil temperature. Root respiration in citrus appears to acclimate rapidly to changes in soil temperature.     

Estimating respiration of roots in soil: Interactions with soil CO2, soil temperature and soil water content

Little information is available on the variability of the dynamics of the actual and observed root respiration rate in relation to abiotic factors. In this study, we describe I) interactions between soil CO₂ concentration, temperature, soil water content and root respiration, and II) the effect of short-term fluctuations of these three environmental factors on the relation between actual and observed root respiration rates. We designed an automated, open, gas-exchange system that allows continuous measurements on 12 chambers with intact roots in soil. By using three distinct chamber designs with each a different path for the air flow, we were able to measure root respiration over a 50-fold range of soil CO₂ concentrations (400 to 25000 ppm) and to separate the effect of irrigation on observed vs. actual root respiration rate. All respiration measurements were made on one-year-old citrus seedlings in sterilized sandy soil with minimal organic material.Root respiration was strongly affected by diurnal fluctuations in temperature (Q₁₀ = 2), which agrees well with the literature. In contrast to earlier findings for Douglas-fir (Qi et al., 1994), root respiration rates of citrus were not affected by soil CO₂ concentrations (400 to 25000 ppm CO₂; pH around 6). Soil CO₂ was strongly affected by soil water content but not by respiration measurements, unless the air flow for root respiration measurements was directed through the soil. The latter method of measuring root respiration reduced soil CO₂ concentration to that of incoming air. Irrigation caused a temporary reduction in CO₂ diffusion, decreasing the observed respiration rates obtained by techniques that depended on diffusion. This apparent drop in respiration rate did not occur if the air flow was directed through the soil. Our dynamic data are used to indicate the optimal method of measuring root respiration in soil, in relation to the objectives and limitations of the experimental conditions.     

Annual variation in soil respiration and its components in a coppice oak forest in Central Italy

In order to investigate the annual variation of soil respiration and its components in relation to seasonal changes in soil temperature and soil moisture in a Mediterranean mixed oak forest ecosystem, we set up a series of experimental treatments in May 1999 where litter (no litter), roots (no roots, by trenching) or both were excluded from plots of 4 m². Subsequently, we measured soil respiration, soil temperature and soil moisture in each plot over a year after the forest was coppiced. The treatments did not significantly affect soil temperature or soil moisture measured over 0–10 cm depth. Soil respiration varied markedly during the year with high rates in spring and autumn and low rates in summer, coinciding with summer drought, and in winter, with the lowest temperatures. Very high respiration rates, however, were observed during the summer immediately after rainfall events. The mean annual rate of soil respiration was 2.9 µ mol m^?2 s^?1, ranging from 1.35 to 7.03 µmol m^?2 s^?1. Soil respiration was highly correlated with temperature during winter and during spring and autumn whenever volumetric soil water content was above 20%. Below this threshold value, there was no correlation between soil respiration and soil temperature, but soil moisture was a good predictor of soil respiration. A simple empirical model that predicted soil respiration during the year, using both soil temperature and soil moisture accounted for more than 91% of the observed annual variation in soil respiration. All the components of soil respiration followed a similar seasonal trend and were affected by summer drought. The Q₁₀ value for soil respiration was 2.32, which is in agreement with other studies in forest ecosystems. However, we found a Q₁₀ value for root respiration of 2.20, which is lower than recent values reported for forest sites. The fact that the seasonal variation in root growth with temperature in Mediterranean ecosystems differs from that in temperate regions may explain this difference. In temperate regions, increases in size of root populations during the growing season, coinciding with high temperatures, may yield higher apparent Q₁₀ values than in Mediterranean regions where root growth is suppressed by summer drought. The decomposition of organic matter and belowground litter were the major components of soil respiration, accounting for almost 55% of the total soil respiration flux. This proportion is higher than has been reported for mature boreal and temperate forest and is probably the result of a short‐term C loss following recent logging at the site. The relationship proposed for soil respiration with soil temperature and soil moisture is useful for understanding and predicting potential changes in Mediterranean forest ecosystems in response to forest management and climate change.     

Photosynthesis,respiration, and carbon allocation of two cool-season perennial grasses in response to surface soil drying

The study was conducted to investigate carbon metabolic responses to surface soil drying for cool-season grasses. Kentucky bluegrass (Poa pratensis L.) and tall fescue (Festuca arundinaceae Schreb.) were grown in a greenhouse in split tubes consisting of two sections. Plants were subjected to three soil moisture regimes: (1) well-watered control; (2) drying of upper 20-cm soil (upper drying); and (3) drying of whole 40-cm soil profile (full drying). Upper drying for 30 d had no dramatic effects on leaf water potential (Ψ_leaf) and canopy photosynthetic rate (P_n) in either grass species compared to the well-watered control, but it reduced canopy respiration rate (R_canopy) and root respiration rate in the top 20 cm of soil (R_top). For both species in the lower 20 cm of wet soil, root respiration rates (R_bottom) were similar to the control levels, and carbon allocation to roots increased with the upper soil drying, particularly for tall fescue. The proportion of roots decreased in the 0-20 cm drying soil, but increased in the lower 20 cm wet soil for both grass species; the increase was greater for tall fescue. The Ψ_leaf, P_n, R_canopy, R_top, R_bottom, and carbon allocation to roots in both soil layers were all significantly higher for upper dried plants than for fully dried plants of both grass species. The reductions in R_canopy and R_top in surface drying soil and increases in root respiration and carbon allocation to roots in lower wet soil could help these grasses cope with surface-soil drought stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

On the assessment of root and soil respiration for soils of different textures: interactions with soil moisture contents and soil CO2 concentrations

Estimates of root and soil respiration are becoming increasingly important in agricultural and ecological research, but there is little understanding how soil texture and water content may affect these estimates. We examined the effects of soil texture on (i) estimated rates of root and soil respiration and (ii) soil CO₂ concentrations, during cycles of soil wetting and drying in the citrus rootstock, Volkamer lemon (Citrus volkameriana Tan. and Pasq.). Plants were grown in soil columns filled with three different soil mixtures varying in their sand, silt and clay content. Root and soil respiration rates, soil water content, plant water uptake and soil CO₂ concentrations were measured and dynamic relationships among these variables were developed for each soil texture treatment. We found that although the different soil textures differed in their plant-soil water relations characteristics, plant growth was only slightly affected. Root and soil respiration rates were similar under most soil moisture conditions for soils varying widely in percentages of sand, silt and clay. Only following irrigation did CO₂ efflux from the soil surface vary among soils. That is, efflux of CO₂ from the soil surface was much more restricted after watering (therefore rendering any respiration measurements inaccurate) in finer textured soils than in sandy soils because of reduced porosity in the finer textured soils. Accordingly, CO₂ reached and maintained the highest concentrations in finer textured soils (> 40 mmol CO₂ mol⁻¹). This study revealed that changes in soil moisture can affect interpretations of root and soil measurements based on CO₂ efflux, particularly in fine textured soils. The implications of the present findings for field soil CO₂ flux measurements are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Elevated growth temperatures reduce the carbon gain of black spruce [Picea mariana (Mill.) B.S.P.]

We explored the effect of high‐growth temperatures on a dominant North American boreal tree, black spruce [Picea mariana (Mill.) B.S.P.]. In 2004 and 2005, we grew black spruce at either 22 °C/16 °C day/night temperatures [low temperature (LT)] or 30°/24 °C [high temperature (HT)] and determined how temperature affected growth, leaf morphology, photosynthesis, respiration and thermotolerance. HT spruce were 20% shorter, 58% lighter, and had a 58% lower root : shoot ratio than LT trees. Mortality was negligible in the LT treatment, but up to 14% of HT seedlings died by the end of the growing season. HT seedlings had a higher photosynthetic temperature optimum, but net photosynthesis at growth temperatures was 19–35% lower in HT than LT trees. HT seedlings had both a lower apparent maximum ribulose‐1,5‐bisphosphate carboxylation capacity (V_cmax) and a lower apparent maximum electron transport rate (J_max) than LT trees, indicating reduced allocation to photosynthetic components. Consistently, HT needles had 26% lower leaf nitrogen content than LT needles. At each measurement temperature, HT seedlings had 20–25% lower respiration rates than LT trees; however, this did not compensate for reduced photosynthetic rates at growth temperature, leading to a greater ratio of dark respiration to net carbon dioxide assimilation rate in HT trees. HT needles had 16% lower concentrations of soluble sugars than LT needles, but similar starch content. Growth at high temperatures increased the thermotolerance of black spruce. HT trees showed less PSII inhibition than LT seedlings and no increase in electrolyte leakage when briefly exposed to 40–57 °C. While trees that develop at high temperatures have enhanced tolerance for brief, extreme heat events, the reduction in root allocation indicates that seedlings will be more susceptible to episodic soil drying and less competitive for belowground resources in future climates of the boreal region.     

Midday depression in root respiration of <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis>: its implication for estimating carbon cycling in forest ecosystems

We observed the phenomenon of midday depression in the rate of tree root respiration. Diurnal changes in the root respiration rate of Quercus crispula and Chamaecyparis obtusa were measured under intact conditions using a closed chamber method and a soil respiration measurement system (LI-6400 with a root respiration chamber) in a forest in the foothills of Mt. Fuji. After the measurement of intact root respiration in the field, the root was excised and taken to the laboratory, and the temperature dependence on the respiration rate of the detached root was measured using an open-flow gas exchange system with an infrared gas analyzer (LI-6252). The measurement was conducted in September 2003, August and November 2005, and June 2006. Whereas the root respiration rate of both species under intact conditions increased with increasing soil and root temperatures from dawn to early morning, the respiration rate decreased around midday from 10:00 to 15:00 despite an increment of soil and root temperatures. There was no clear relationship between the intact root respiration rate and root temperature in either species, although the detached root respiration rate of both increased exponentially with the temperature. The amount of the CO₂ efflux estimated using the temperature dependence of detached root respiration tended to underestimate the actual measurement value (intact respiration rate) by 20–50% in both species. These results indicate that evaluating midday depression in root respiration would be important for a more accurate estimation of the carbon cycle or net ecosystem production in forests.     

Inverse determination of heterotrophic soil respiration response to temperature and water content under field conditions

Heterotrophic soil respiration is an important flux within the global carbon cycle. Exact knowledge of the response functions for soil temperature and soil water content is crucial for a reliable prediction of soil carbon turnover. The classical statistical approach for the in situ determination of the temperature response (Q₁₀ or activation energy) of field soil respiration has been criticised for neglecting confounding factors, such as spatial and temporal changes in soil water content and soil organic matter. The aim of this paper is to evaluate an alternative method to estimate the temperature and soil water content response of heterotrophic soil respiration. The new method relies on inverse parameter estimation using a 1-dimensional CO₂ transport and carbon turnover model. Inversion results showed that different formulations of the temperature response function resulted in estimated response factors that hardly deviated over the entire range of soil water content and for temperature below 25°C. For higher temperatures, the temperature response was highly uncertain due to the infrequent occurrence of soil temperatures above 25°C. The temperature sensitivity obtained using inverse modelling was within the range of temperature sensitivities estimated from statistical processing of the data. It was concluded that inverse parameter estimation is a promising tool for the determination of the temperature and soil water content response of soil respiration. Future synthetic model studies should investigate to what extent the inverse modelling approach can disentangle confounding factors that typically affect statistical estimates of the sensitivity of soil respiration to temperature and soil water content.     

Physiological Responses of Apple Trees to Supraoptimal Root Temperature

Ungrafted apple rootstocks were grown in sand cultures at constant root temperatures between 20°C to 40°C. Temperatures of 30°C and above reduced root and shoot growth. Serious damage to the leaves occurred at 35°C and above. The O₂ consumption, CO₂ evolution and respiratory quotient (RQ) of the roots showed maximum values at 35°C. Different rootstock cultivars varied greatly in their susceptibility to damage by supraoptimal root temperatures apparently due to anaerobic respiration. The more susceptible ones differed from resistant types in the larger amount of ethanol they accumulated in their roots at supraoptimal root temperature, and the more severe reduction in the malic acid content of the roots at such temperature. Acetaldehyde was also found in roots and leaves at supraoptimal root temperatures, whereas the organic acid content of the leaves tended to decrease. Supraoptimal root temperature also caused a reduction of cytokinins in both roots and leaves accompanied by a reduction in the leaf chlorophyll content. This could be prevented by the application of kinetin or benzyladenine to the leaves. In a short experiment a rise in root temperature up to 40°C caused an increase in transpiration and a decrease in the resistance of the leaves to the passage of water vapor, whereas in prolonged experiments transpiration reached a maximum and leaf resistance a minimum at 30°C. The leaf water potential increased also with increasing root temperature. Leaf temperature increased with increasing root temperature, irrespective of increasing or decreasing transpiration rates.     

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range,Alaska, USA

Summary Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO₂ m^–2 day^–1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO₂ efflux in this habitat. In parallel with the field measurements of CO₂ efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO₂ efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO₂ efflux and carbon balance.     

Respiration activity of ectomycorrhizas from Cenococcum geophilum and Lactarius sp. in relation to soil water potential in five beech forests

Forest trees are involved in root symbioses with hundreds of species of ectomycorrhizal fungi which constitute functional guilds able to improve the water and mineral nutrition of host trees. In temperate ecosystems, water shortage is a main factor limiting tree vitality. To assess how soil water conditions affected the physiological state of beech (Fagus silvatica L.) ectomycorrhizal roots, we monitored glucose respiration of two ectomycorrhizal types (Lactarius sp. and Cenococcum geophilum) during two complete growing seasons. Five stands of contrasting soil conditions were chosen in north-eastern France. The top soil horizons were equipped with micropsychrometers for measuring water potential and temperature. Glucose respiration on individual ectomycorrhizas was measured in vitro by trapping [¹⁴C]-CO₂ from radiolabelled glucose. For soil water potential <-0.2 MPa, the potential respiration activity of C. geophilumectomycorrhizas was significantly less altered than that of Lactariussp. ectomycorrhizas, indicating that C. geophilumis more likely than Lactariussp. to maintain the physiological integrity of beech roots facing drought stress.     

Tree root and soil heterotrophic respiration as revealed by girdling of boreal Scots pine forest: extending observations beyond the first year

Limitations in available techniques to separate autotrophic (root) and soil heterotrophic respiration have hampered the understanding of forest C cycling. The former is here defined as respiration by roots, their associated mycorrhizal fungi and other micro‐organisms in the rhizosphere directly dependent on labile C compounds leaked from roots. In order to separate the autotrophic and heterotrophic components of soil respiration, all Scots pine trees in 900 m² plots were girdled to instantaneously terminate the supply of current photosynthates from the tree canopy to roots. Högberg et al. (Nature 411, 789–792, 2001) reported that autotrophic activity contributed up to 56% of total soil respiration during the first summer of this experiment. They also found that mobilization of stored starch (and likely also sugars) in roots after girdling caused an increased apparent heterotrophic respiration on girdled plots. Herein a transient increase in the δ¹³C of soil CO₂ efflux after girdling, thought to be due to decomposition of ¹³C‐enriched ectomycorrhizal mycelium and root starch and sugar reserves, is reported. In the second year after girdling, when starch reserves of girdled tree roots were exhausted, calculated root respiration increased up to 65% of total soil CO₂ efflux. It is suggested that this estimate of its contribution to soil respiration is more precise than the previous based on one year of observation. Heterotrophic respiration declined in response to a 20‐day‐long 6 °C decline in soil temperature during the second summer, whereas root respiration did not decline. This did not support the idea that root respiration should be more sensitive to variations in soil temperature. It is suggested that above‐ground photosynthetic activity and allocation patterns of recent photosynthates to roots should be considered in models of responses of forest C balances to global climate change.     

Carbon exchange in biological soil crust communities under differential temperatures and soil water contents: implications for global change

Biological soil crusts (biocrusts) are an integral part of the soil system in arid regions worldwide, stabilizing soil surfaces, aiding vascular plant establishment, and are significant sources of ecosystem nitrogen and carbon. Hydration and temperature primarily control ecosystem CO₂ flux in these systems. Using constructed mesocosms for incubations under controlled laboratory conditions, we examined the effect of temperature (5–35 °C) and water content (WC, 20–100%) on CO₂ exchange in light (cyanobacterially dominated) and dark (cyanobacteria/lichen and moss dominated) biocrusts of the cool Colorado Plateau Desert in Utah and the hot Chihuahuan Desert in New Mexico. In light crusts from both Utah and New Mexico, net photosynthesis was highest at temperatures >30 °C. Net photosynthesis in light crusts from Utah was relatively insensitive to changes in soil moisture. In contrast, light crusts from New Mexico tended to exhibit higher rates of net photosynthesis at higher soil moisture. Dark crusts originating from both sites exhibited the greatest net photosynthesis at intermediate soil water content (40–60%). Declines in net photosynthesis were observed in dark crusts with crusts from Utah showing declines at temperatures >25 °C and those originating from New Mexico showing declines at temperatures >35 °C. Maximum net photosynthesis in all crust types from all locations were strongly influenced by offsets in the optimal temperature and water content for gross photosynthesis compared with dark respiration. Gross photosynthesis tended to be maximized at some intermediate value of temperature and water content and dark respiration tended to increase linearly. The results of this study suggest biocrusts are capable of CO₂ exchange under a wide range of conditions. However, significant changes in the magnitude of this exchange should be expected for the temperature and precipitation changes suggested by current climate models.     

Carbon use in root respiration as affected by elevated atmospheric O2

The use of fossil fuel is predicted to cause an increase of the atmospheric CO₂ concentration, which will affect the global pattern of temperature and precipitation. It is therefore essential to incorporate effects of temperature and water supply on the carbon requirement for root respiration of plants to predict effects of elevated [CO₂] on the carbon budget of natural and managed systems.There is insufficient information to support the contentention that an increase in the concentration of CO₂ in the atmosphere will enhance the CO₂ concentration in the soil to an extent that is likely to affect root respiration. Moreover, there is no convincing evidence for a direct effect of elevated atmospheric [CO₂] on the rate of root respiration per unit root mass or the fraction of carbon required for root respiration. However, there are likely to be indirect effects of elevated [CO₂] on the carbon requirement of plants in natural systems.Firstly, it is very likely that the carbon requirement of root respiration relative to that fixed in photosynthesis will increase when elevated [CO₂] induces a decrease in nutrient status of the plants. Although earlier papers have emphasized that elevated [CO₂] favours investment of biomass in roots relative to that in leaves, these are in fact indirect effects. The increase in root weight ratio is due to the more rapid depletion of nutrients in the root environment as a consequence of enhanced growth. This will decrease the specific rate of root respiration, but increase the carbon requirement as a fraction of the carbon fixed in photosynthesis. It is likely that these effects will be minor in systems where the nutrient supply is very high, e.g. in many managed arable systems, and increase with decreasing soil fertility, i.e. in many natural systems.Secondly, a decrease in rainfall in some parts of the world may cause a shortage in water supply which favours the carbon partitioning to roots. Water stress is likely to reduce rates of root respiration per unit root mass, but enhance the fraction of total assimilates required for root respiration, due to greater allocation of biomass to roots.Increased temperatures are unlikely to affect the specific rate of root respiration in all species. Broadly generalized, the effect of temperature on biomass allocation is that the relative investment of biomass in roots is lowest at a certain optimum temperature and increases at both higher and lower temperatures. The root respiration of some species acclimates to growth temperature, so that the effect of global temperature rise is entirely accounted for by the effect of temperature on biomass allocation. The specific rate of root respiration of other species will increase with global warming. In response to global warming the carbon requirement of roots is likely to decrease in temperate regions, when temperatures are suboptimal for the roots' capacity to acquire water. Here global warming will induce a smaller biomass allocation to the roots. Conversely, the carbon requirements are more likely to increase in mediterranean environments, where temperatures are often supraoptimal and a rise in temperature will induce greater allocation of biomass to the roots.     

Soil CO2 concentration does not affect growth or root respiration in bean or citrus

Contrasting effects of soil CO₂ concentration on root respiration rates during short-term CO₂ exposure, and on plant growth during long-term CO₂ exposure, have been reported. Here we examine the effects of both short- and long-term exposure to soil CO₂ on the root respiration of intact plants and on plant growth for bean (Phaseolus vulgaris L.) and citrus (Citrus volkameriana Tan. & Pasq.). For rapidly growing bean plants, the growth and maintenance components of root respiration were separated to determine whether they differ in sensitivity to soil CO₂. Respiration rates of citrus roots were unaffected by the CO₂ concentration used during the respiration measurements (200 and 2000 μmol mol⁻¹), regardless of the soil CO₂, concentration during the previous month (600 and 20 000 μmol mol⁻¹). Bean plants were grown with their roots exposed to either a natural CO₂ diffusion gradient, or to an artificially maintained CO₂ concentration of 600 or 20 000 μmol mol⁻¹. These treatments had no effect on shoot and root growth. Growth respiration and maintenance respiration of bean roots were also unaffected by CO₂ pretreatment and the CO₂ concentration used during the respiration measurements (200–2000 μmol mol⁻¹). We conclude that soil CO₂ concentrations in the range likely to be encountered in natural soils do not affect root respiration in citrus or bean.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Water relations and root activities of Buchloe dactyloides and Zoysia japonica in response to localized soil drying

Effects of localized soil drought stress on water relations, root growth, and nutrient uptake were examined in drought tolerant ‘Prairie’ buffalograss [Buchloe dactyloides (Nutt.) Engelm.] and sensitive ‘Meyer’ zoysiagrass (Zoysia japonica Steud.). Grasses were grown in small rhizotrons in a greenhouse and subjected to three soil moisture regimes: (1) watering the entire 80-cm soil profile (well-watered control); (2) drying 0–40 cm soil and watering the lower 40 cm (partially dried); (3) and drying the entire soil profile (fully dried). Drying the 0–40 cm soil for 28 days had no effect on leaf water potential (Ψ _leaf ) in Prairie buffalograss compared to the well-watered control but reduced that in Meyer zoysiagrass. Root elongation rate was greater for Prairie buffalograss than Meyer zoysiagrass under well-watered or fully dried conditions. Rooting depth increased with surface soil drying; with Prairie buffalograss having a larger proportion of roots in the lower 40 cm than Meyer zoysiagrass. The higher rates of water uptake in the deeper soil profile in the partially dried compared to the well-watered treatment and by Prairie buffalograss compared to Meyer zoysiagrass could be due to differences in root distribution. Root ¹⁵N uptake for Prairie buffalograss was higher in 0–20 cm drying soil in the partially dried treatment than in the fully dried treatment. Diurnal fluctuations in soil water content in the upper 20 cm of soil when the lower 40 cm were well-watered indicated water efflux from the deeper roots to the drying surface soil. This could help sustain root growth, maintain nutrient uptake in the upper drying soil layer, and prolong turfgrass growth under localized drying conditions, especially for the deep-rooted Prairie buffalograss. This revised version was published online in June 2006 with corrections to the Cover Date.     

Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest

Variation in soil temperature can account for most of the seasonal and diel variation in soil CO₂ efflux, but the temperature effect is not always consistent, and other factors such as soil water content are known to influence soil respiration. The objectives of this research were to study the spatial and temporal variation in soil respiration in a temperate forested landscape and to evaluate temperature and soil water functions as predictors of soil respiration. Soil CO₂ fluxes were measured with chambers throughout an annual cycle in six study areas at the Harvard Forest in central Massachusetts that include soil drainage classes from well drained to very poorly drained. The mean annual estimate of soil CO₂ efflux was 7.2 Mg ha^–1, but ranged from 5.3 in the swamp site to 8.5 in a well-drained site, indicating that landscape heterogeneity is related to soil drainage class. An exponential function relating CO₂ fluxes to soil temperature accounted for 80% of the seasonal variation in fluxes across all sites (Q₁₀ = 3.9), but the Q₁₀ ranged from 3.4 to 5.6 for the individual study sites. A significant drought in 1995 caused rapid declines in soil respiration rates in August and September in five of the six sites (a swamp site was the exception). This decline in CO₂ fluxes correlated exponentially with decreasing soil matric potential, indicating a mechanistic effect of drought stress. At moderate to high water contents, however, soil water content was negatively correlated with soil temperature, which precluded distinguishing between the effects of these two confounded factors on CO₂ flux. Occurrence of high Q₁₀ values and variation in Q₁₀ values among sites may be related to: (i) confounding effects of high soil water content; (ii) seasonal and diel patterns in root respiration and turnover of fine roots that are linked to above ground phenology and metabolism; and (iii) variation in the depth where CO₂ is produced. The Q₁₀ function can yield reasonably good predictions of annual fluxes of CO₂, but it is a simplification that masks responses of root and microbial processes to variation in temperature and water content throughout the soil.     

降雨对旱作春玉米农田土壤呼吸动态的影响

土壤呼吸是调控全球碳平衡和气候变化的关键过程之一,降雨作为重要的扰动因子,在不同区域和不同环境条件下,对土壤呼吸具有复杂的影响.研究降雨对农田土壤呼吸及其分量的影响,对准确预测未来气候变化下陆地生态系统碳平衡具有重要意义.对黄土高原东部典型春玉米农田生态系统生长季内3次降雨前后土壤呼吸及其分量进行了原位连续观测,结果表明:在土壤湿润的条件下,降雨对春玉米农田土壤呼吸及其分量具有明显的抑制作用,在土壤湿度大于27％后土壤呼吸及其分量随土壤湿度上升呈明显下降,且对温度的敏感性降低.土壤呼吸及其分量在降雨前后的变化受土壤温度和土壤湿度的共同影响.降雨量、降雨历时和雨前土壤含水量决定了土壤呼吸及其分量对降雨响应的程度和时长.土壤呼吸及其分量对土壤温度的敏感性各不相同,微生物呼吸对温度的敏感性最高,Q10为5.14;其次是土壤呼吸,Q10为3.86;根呼吸的温度敏感性相对最低,Q10为3.24.由于土壤呼吸分量对温度和湿度的敏感性不同,降雨后根呼吸的比例有所升高.