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.     

Effects of shoot excision on in situ soil and root respiration of wheat and soybean under drought stress

Little information is available about the variability of root-derived respiration rate in relation to biotic factors such as photosynthesis and substrate availability in roots. Here we examine the role of decreased carbohydrates availability on root-derived respiration through removal of above ground biomass. Spring wheat (Triticum aestivum L. cv. Longchun 8139) and soybean (Glycina max L. cv. Tianchan 2) were grown in the field under a moveable rain shelter, and subjected to three different water regimes: (1) well-watered control; (2) moderate drought stress, and (3) severe drought stress. Root-derived respiration before and after shoot clipping, and the concentration of total nonstructural carbohydrate, malic and citric acid were measured for spring wheat and soybean. Root-derived CO₂ flux and total nonstructural carbohydrate concentration of clipped wheat decreased by 38% and 31%, respectively. However, for soybean the root- derived CO₂ flux and total nonstructural carbohydrate concentrations were only 58% and 62% of control, respectively, indicating the root respiration rate was controlled by the availability of carbon in the root. A significant positive correlation between total nonstructural carbohydrate concentration of the root and soil water content was observed in unclipped plants. Total nonstructural carbohydrate contributed 93% of the variance in root-derived respiration. Our results clearly show, that in the field, the availability of carbon substrate in roots determines root-derived respiration and plays a key link between soil moisture and root-derived respiration. A period of time is needed for root respiration to return to “steady-state” after shoot removal and this period needed is strongly dependent on species and soil water content.     

Forest soil respiration rate and δ13C is regulated by recent above ground weather conditions

Soil respiration, a key component of the global carbon cycle, is a major source of uncertainty when estimating terrestrial carbon budgets at ecosystem and higher levels. Rates of soil and root respiration are assumed to be dependent on soil temperature and soil moisture yet these factors often barely explain half the seasonal variation in soil respiration. We here found that soil moisture (range 16.5–27.6% of dry weight) and soil temperature (range 8–17.5°C) together explained 55% of the variance (cross-validated explained variance; Q²) in soil respiration rate (range 1.0–3.4 mol C m^–2 s^–1) in a Norway spruce (Picea abies) forest. We hypothesised that this was due to that the two components of soil respiration, root respiration and decomposition, are governed by different factors. We therefore applied PLS (partial least squares regression) multivariate modelling in which we, together with below ground temperature and soil moisture, used the recent above ground air temperature and air humidity (vapour pressure deficit, VPD) conditions as x-variables. We found that air temperature and VPD data collected 1–4 days before respiration measurements explained 86% of the seasonal variation in the rate of soil respiration. The addition of soil moisture and soil temperature to the PLS-models increased the Q² to 93%. ¹³C analysis of soil respiration supported the hypotheses that there was a fast flux of photosynthates to root respiration and a dependence on recent above ground weather conditions. Taken together, our results suggest that shoot activities the preceding 1–6 days influence, to a large degree, the rate of root and soil respiration. We propose this above ground influence on soil respiration to be proportionally largest in the middle of the growing season and in situations when there is large day-to-day shifts in the above ground weather conditions. During such conditions soil temperature may not exert the major control on root respiration.     

Root production and turnover in an upland grassland subjected to artificial soil warming respond to radiation flux and nutrients, not temperature

Root demographic processes (birth and death) were measured using minirhizotrons in the soil warming experiments at the summit of Great Dun Fell, United Kingdom (845 m). The soil warming treatment raised soil temperature at 2 cm depth by nearly 3°C. The first experiment ran for 6 months (1994), the second for 18 (1995–1996). In both experiments, heating increased death rates for roots, but birth rates were not significantly increased in the first experiment. The lack of stimulation of death rate in 1996 is probably an artefact, caused by completion of measurements in late summer of 1996, before the seasonal demography was concluded: root death continued over the winter of 1995–1996. Measurements of instantaneous death rates confirmed this: they were accelerated by warming in the second experiment. In the one complete year (1995–1996) in which measurements were taken, net root numbers by the end of the year were not affected by soil warming. The best explanatory environmental variable for root birth rate in both experiments was photosynthetically active radiation (PAR) flux, averaged over the previous 5 (first experiment) or 10 days (second experiment). In the second experiment, the relationship between birth rate and PAR flux was steeper and stronger in heated than in unheated plots. Death rate was best explained by vegetation temperature. These results provide further evidence that root production acclimates to temperature and is driven by the availability of photosynthate. The stimulation of root growth due to soil warming was almost certainly the result of changes in nutrient availability following enhanced decomposition. Received: 4 May 1999 / Accepted: 18 May 1999     

Representative estimates of soil and ecosystem respiration in an old beech forest

Respiration has been proposed to be the main determinant of the carbon balance in European forests and is thus essential for our understanding of the carbon cycle. However, the choice of experimental design strongly affects estimates of annual respiration and of the contribution of soil respiration to total ecosystem respiration. In a detailed study of ecosystem and soil respiration fluxes in an old unmanaged deciduous forest in Central Germany over 3 years (2000–2002), we combined soil chamber and eddy covariance measurements to obtain a comprehensive picture of respiration in this forest. The closed portable chambers offered to investigate spatial variability of soil respiration and its controls while the eddy covariance system offered continuous measurements of ecosystem respiration. Over the year, both fluxes were mainly correlated with temperature. However, when soil moisture sank below 23 vol.% in the upper 6 cm, water limitations also became apparent. The temporal resolution of the eddy covariance system revealed that relatively high respiration rates occurred during budbreak due to increased metabolic activity and after leaf fall because of increased decomposition. Spatial variability in soil respiration rates was large and correlated with fine root biomass (r ² = 0.56) resulting in estimates of annual efflux varying across plots from 730 to 1,258 (mean 898) g C m⁻² year⁻¹. Power function calculations showed that achieving a precision in the soil respiration estimate of 20% of the full population mean at a confidence level of 95%, requires about eight sampling locations. Our results can be used as guidelines to improve the representativeness of soil respiration measurements by nested sampling designs, being applied in long-term and large-scale carbon sequestration projects such as FLUXNET and CarboEurope.     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.     

Main determinants of forest soil respiration along an elevation/temperature gradient in the Italian Alps

The main determinants of soil respiration were investigated in 11 forest types distributed along an altitudinal and thermal gradient in the southern Italian Alps (altitudinal range 1520 m, range in mean annual temperature 7.8°C). Soil respiration, soil carbon content and principal stand characteristics were measured with standardized methods. Soil CO₂ fluxes were measured at each site every 15–20 days with a closed dynamic system (LI‐COR 6400) using soil collars from spring 2000 to spring 2002. At the same time, soil temperature at a depth of 10 cm and soil water content (m³ m^?3) were measured at each collar. Soil samples were collected to a depth of 30 cm and stones, root content and bulk density were determined in order to obtain reliable estimates of carbon content per unit area (kg C m^?2). Soil respiration and temperature data were fitted with a simple logistic model separately for each site, so that base respiration rates and mean annual soil respiration were estimated. Then the same regression model was applied to all sites simultaneously, with each model parameter being expressed as a linear function of site variables. The general model explained about 86% of the intersite variability of soil respiration. In particular, soil mean annual temperature explained the most of the variance of the model (0.41), followed by soil temperature interquartlile range (0.24), soil carbon content (0.16) and soil water content (0.05).     

Phosphate Addition and Plant Species Alters Microbial Community Structure in Acidic Upland Grassland Soil

Agricultural improvement (addition of fertilizers, liming) of seminatural acidic grasslands across Ireland and the UK has resulted in significant shifts in floristic composition, soil chemistry, and microbial community structure. Although several factors have been proposed as responsible for driving shifts in microbial communities, the exact causes of such changes are not well defined. Phosphate was added to grassland microcosms to investigate the effect on fungal and bacterial communities. Plant species typical of unimproved grasslands (Agrostis capillaris, Festuca ovina) and agriculturally improved grasslands (Lolium perenne) were grown, and phosphate was added 25 days after seed germination, with harvesting after a further 50 days. Phosphate addition significantly increased root biomass (p < 0.001) and shoot biomass (p < 0.05), soil pH (by 0.1 U), and microbial activity (by 5.33 mg triphenylformazan [TPF] g⁻¹ soil; p < 0.001). A slight decrease (by 0.257 mg biomass-C g⁻¹ soil; p < 0.05) in microbial biomass after phosphate addition was found. The presence of plant species significantly decreased soil pH (p < 0.05; by up to 0.2 U) and increased microbial activity (by up to 6.02 mg TPF g⁻¹ soil) but had no significant effect on microbial biomass. Microbial communities were profiled using automated ribosomal intergenic spacer analysis. Multidimensional scaling plots and canonical correspondence analysis revealed that phosphate addition and its interactions with upland grassland plant species resulted in considerable changes in the fungal and bacterial communities of upland soil. The fungal community structure was significantly affected by both phosphate (R = 0.948) and plant species (R = 0.857), and the bacterial community structure was also significantly affected by phosphate (R = 0.758) and plant species (R = 0.753). Differences in microbial community structure following P addition were also revealed by similarity percentage analysis. These data suggest that phosphate application may be an important contributor to microbial community structural change during agricultural management of upland grasslands.     

Effects of temperature on photosynthesis of two morphologically contrasting plant species along an altitudinal gradient in the tropical high Andes

The effects of temperature on photosynthesis of a rosette plant growing at ground level, Acaena cylindrostachya R. et P., and an herb that grows 20–50 cm above ground level, Senecio formosus H.B.K., were studied along an altitudinal gradient in the Venezuelan Andes. These species were chosen in order to determine – in the field and in the laboratory – how differences in leaf temperature, determined by plant form and microenvironmental conditions, affect their photosynthetic capacity. CO₂ assimilation rates (A) for both species decreased with increasing altitude. For Acaena leaves at 2900 m, A reached maximum values above 9 μmol m⁻² s⁻¹, nearly twice as high as maximum A found at 3550 m (5.2) or at 4200 m (3.9). For Senecio leaves, maximum rates of CO₂ uptake were 7.5, 5.8 and 3.6 μmol m⁻² s⁻¹ for plants at 2900, 3550 and 4200 m, respectively. Net photosynthesis-leaf temperature relations showed differences in optimum temperature for photosynthesis (A _o.t.) for both species along the altitudinal gradient. Acaena showed similar A _o.t. for the two lower altitudes, with 19.1°C at 2900 m and 19.6°C at 3550 m, while it increased to 21.7°C at 4200 m. Maximum A for this species at each altitude was similar, between 5.5 and 6.0 μmol m⁻² s⁻¹. For the taller Senecio, A _o.t. was more closely related to air temperatures and decreased from 21.7°C at 2900 m, to 19.7°C at 3550 m and 15.5°C at 4200 m. In this species, maximum A was lower with increasing altitude (from 6.0 at 2900 m to 3.5 μmol m⁻² s⁻¹ at 4200 m). High temperature compensation points for Acaena were similar at the three altitudes, c. 35°C, but varied in Senecio from 37°C at 2900 m, to 39°C at 3550 m and 28°C at 4200 m. Our results show how photosynthetic characteristics change along the altitudinal gradient for two morphologically contrasting species influenced by soil or air temperatures. Received: 5 July 1997 / Accepted: 25 October 1997     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

Flooding tolerance of Carex species. II. Root gas-exchange capacity

Root CO₂ and O₂ gas exchange were measured in young Carex extensa Good. (flooding sensitive), C. remota L. and C. pseudocyperus L. (both flooding tolerant) plants, precultured either aerobically or anaerobically. Temperature changes form 21 to 11 °C had small effects on root CO₂ release from respiration. In C. extensa, root respiration rates decreased when plants were precultured anaerobically, while in C. pseudocyperus preculture conditions had no effect on root CO₂ release. In contrast to CO₂, temperature decrease significantly enhanced radial oxygen loss from the roots during the light phase, indicating that at 20 °C the O₂ transported form the shoot to the root met the demand for root respiration quite well, while at 11 °C excess O₂ entered the root and was released into the anaerobic nutrient solution. In C. remota and C. pseudocyperus, the maximal O₂ concentration of a previously anaerobic nutrient solution was attained after several days of equilibration with the atmosphere through the plant body and was approximately one-third of that found in C. extensa, indicating that the diffusion resistance of the root/rhizosphere interface to O₂ is much lower in the C. extensa root than in the flooding-tolerant Carex species. A calculation of the maximal attainable root length that can be sustained by pure O₂ diffusion from CO₂ exchange, and anatomical data obtained earlier, revealed that longitudinal diffusion of O₂ through the root is sufficient for the oxygen supply of the root. It is concluded that the postulate of a gas mass-flow into the root is not necessary for the understanding of flooding tolerance of Carex species. Received: 10 February 1998 / Accepted: 2 July 1998     

Dynamics of soil respiration in sparse <Emphasis Type="Italic">Ulmus pumila</Emphasis> woodland under semi-arid climate

Sparse Ulmus pumila woodlands play an important role in contributing to ecosystem function in semi-arid grassland of northern China. To understand the key attributes of soil carbon cycling in U. pumila woodland, we studied dynamics of soil respiration in the canopy field (i.e., the projected crown cover area) and the open field at locations differing in distance (i.e., at 1–1.5, 3–4, 10, and >15 m) to tree stems from July through September of 2005, and measured soil biotic factors (e.g., fine root mass, soil microbial biomass, and activity) and abiotic factors [e.g., soil water content (SWC) and organic carbon] in mid-August. Soil respiration was further separated into root component and microbial component at the end of the field measurement in September. Results showed that soil respiration had a significant exponent relationship with soil temperature at 10-cm depth. The temperature sensitivity index of soil respiration, Q ₁₀, was lower than the global average of 2.0, and declined significantly (P < 0.05) with distance. The rate of soil respiration was generally greater in the canopy field than in the open field; monthly mean of soil respiration was 305.5–730.8 mg CO₂ m⁻² h⁻¹ in the canopy field and 299.6–443.1 mg CO₂ m⁻² h⁻¹ in the open field from July through September; basal soil respiration at 10°C declined with distance, and varied from ~250 mg CO₂ m⁻² h⁻¹ near tree stems to <200 mg CO₂ m⁻² h⁻¹ in the open field. Variations in soil respiration with distance were consistent with patterns of SWC, fine root mass, microbial biomass and activities. Regression analysis indicated that soil respiration was tightly coupled with microbial respiration and only weakly related to root respiration. Overall, variations in SWC, soil nutrients, microbial biomass, and microbial activity are largely responsible for the spatial heterogeneity of soil respiration in this semi-arid U. pumila woodland.     

Root Respiration,Photosynthesis and Grain Yield of Two Spring Wheat in Response to Soil Drying

The effects of soil water regime and wheat cultivar, differing in drought tolerance with respect to root respiration and grain yield, were investigated in a greenhouse experiment. Two spring wheat (Triticum aestivum) cultivars, a drought sensitive (Longchun 8139-2) and drought tolerant (Dingxi 24) were grown in PVC tubes (120 cm in length and 10 cm in diameter) under an automatic rain-shelter. Plants were subjected to three soil moisture regimes: (1) well-watered control (85% field water capacity, FWC); (2) moderate drought stress (50% FWC) and (3) severe drought stress (30% FWC). The aim was to study the influence of root respiration on grain yield under soil drying conditions. In the experiment, severe drought stress significantly (p < 0.05) reduced shoot and root biomass, photosynthesis and root respiration rate for both cultivars, but the extent of the decreases was greater for Dingxi 24 compared to that for Longchun 8139-2. Compared with Dingxi 24, 0.04 and 0.07 mg glucose m⁻² s⁻¹ of additional energy, equivalent to 0.78 and 1.43 J m⁻² s⁻¹, was used for water absorption by Longchun 8139-2 under moderate and severe drought stress, respectively. Although the grain yield of both cultivars decreased with declining soil moisture, loss was greater in Longchun 8139-2 than in Dingxi 24, especially under severe drought stress. The drought tolerance cultivar (Dingxi 24), had a higher biomass and metabolic activity under severe drought stress compared to the sensitive cultivar (Longchun 8139-2), which resulted in further limitation of grain yield. Results show that root respiration, carbohydrates allocation (root:shoot ratio) and grain yield were closely related to soil water status and wheat cultivar. Reductions in root respiration and root biomass under severe soil drying can improve drought tolerant wheat growth and physiological activity during soil drying and improve grain yield, and hence should be advantageous over a drought sensitive cultivar in arid regions.     

Carbon balance and allocation of assimilated CO2 in Scots pine, Norway spruce, and Silver birch seedlings determined with gas exchange measurements and 14C pulse labelling

Carbon dioxide is released from the soil to the atmosphere in heterotrophic respiration when the dead organic matter is used for substrates for soil micro-organisms and soil animals. Respiration of roots and mycorrhiza is another major source of carbon dioxide in soil CO₂ efflux. The partitioning of these two fluxes is essential for understanding the carbon balance of forest ecosystems and for modelling the carbon cycle within these ecosystems. In this study, we determined the carbon balance of three common tree species in boreal forest zone, Scots pine, Norway spruce, and Silver birch with gas exchange measurements conducted in laboratory in controlled temperature and light conditions. We also studied the allocation pattern of assimilated carbon with ¹⁴C pulse labelling experiment. The photosynthetic light responses of the tree species were substantially different. The maximum photosynthetic capacity (P _max) was 2.21 μg CO₂ s⁻¹ g⁻¹ in Scots pine, 1.22 μg CO₂ s⁻¹ g⁻¹ in Norway spruce and 3.01 μg CO₂ s⁻¹ g⁻¹ in Silver birch seedlings. According to the pulse labelling experiments, 43–75% of the assimilated carbon remained in the aboveground parts of the seedlings. The amount of carbon allocated to root and rhizosphere respiration was about 9–26%, and the amount of carbon allocated to root and ectomycorrhizal biomass about 13–21% of the total assimilated CO₂. The ¹⁴CO₂ pulse reached the root system within few hours after the labelling and most of the pulse had passed the root system after 48 h. The transport rate of carbon from shoot to roots was fastest in Silver birch seedlings.     

Physiological adaptations to phosphorus deficiency during proteoid root development in white lupin

Release of large amounts of citric acid from specialized root clusters (proteoid roots) of phosphorus (P)-deficient white lupin (Lupinus albus L.) is an efficient strategy for chemical mobilization of sparingly available P sources in the rhizosphere. The present study demonstrates that increased accumulation and exudation of citric acid and a concomitant release of protons were predominantly restricted to mature root clusters in the later stages of P deficiency. Inhibition of citrate exudation by exogenous application of anion-channel blockers such as ethacrynic- and anthracene-9-carboxylic acids may indicate involvement of an anion channel. Phosphorus-deficiency-induced accumulation and subsequent exudation of citric acid seem to be a consequence of both increased biosynthesis and reduced metabolization of citric acid in the proteoid root tissue, indicated by increased in-vitro activity and enzyme protein levels of phosphoenolpyruvate carboxylase (EC 4.1.1.31), and reduced activity of aconitase (EC 4.2.1.3) and root respiration. Similar to citric acid, acid phosphatase, which is secreted by roots and involved in the mobilization of the organic soil P fraction, was released predominantly from proteoid roots of P-deficient plants. Also ³³Pi uptake per unit root fresh-weight was increased by approximately 50% in juvenile and mature proteoid root clusters compared to apical segments of non-proteoid roots. Kinetic studies revealed a K _m of 30.7 μM for Pi uptake of non-proteoid root apices in P-sufficient plants, versus K _m values of 8.5–8.6 μM for non-proteoid and juvenile proteoid roots under P-deficient conditions, suggesting the induction of a high-affinity Pi-uptake system. Obviously, P-deficiency-induced adaptations of white lupin, involved in P acquisition and mobilization of sparingly available P sources, are predominantly confined to proteoid roots, and moreover to distinct stages during proteoid root development. Received: 10 September 1998 / Accepted: 22 December 1998     

Carbon dynamics and budget in a Zoysia japonica grassland, central Japan

The ecosystem carbon budget was estimated in a Japanese Zoysia japonica grassland. The green biomass started to grow in May and peaked from mid-July to September. Seasonal variations in soil CO₂ flux and root respiration were mediated by changes in soil temperature. Annual soil CO₂ flux was 1,121.4 and 1,213.6 g C m⁻² and root respiration was 471.0 and 544.3 g C m⁻² in 2007 and 2008, respectively. The root respiration contribution to soil CO₂ flux ranged from 33% to 71%. During the growing season, net primary production (NPP) was 747.5 and 770.1 g C m⁻² in 2007 and 2008, respectively. The biomass removed by livestock grazing (GL) was 122.1 and 102.7 g C m⁻², and the livestock returned 28.2 and 25.6 g C m⁻² as fecal input (FI) in 2007 and 2008, respectively. The decomposition of FI (DL, the dry weight loss due to decomposition) was very low, 1.5 and 1.4 g C m⁻², in 2007 and 2008. Based on the values of annual NPP, soil CO₂ flux, root respiration, GL, FI, and DL, the estimated carbon budget of the grassland was 1.7 and 22.3 g C m⁻² in 2007 and 2008, respectively. Thus, the carbon budget of this Z. japonica grassland ecosystem remained in equilibrium with the atmosphere under current grazing conditions over the 2 years of the study.     

Stem respiration of ponderosa pines grown in contrasting climates: implications for global climate change

We examined the effects of climate and allocation patterns on stem respiration in ponderosa pine (Pinus ponderosa) growing on identical substrate in the cool, moist Sierra Nevada mountains and the warm, dry, Great Basin Desert. These environments are representative of current climatic conditions and those predicted to accompany a doubling of atmospheric CO₂, respectively, throughout the range of many western north American conifers. A previous study found that trees growing in the desert allocate proportionally more biomass to sapwood and less to leaf area than montane trees. We tested the hypothesis that respiration rates of sapwood are lower in desert trees than in montane trees due to reduced stem maintenance respiration (physiological acclimation) or reduced construction cost of stem tissue (structural acclimation). Maintenance respiration per unit sapwood volume at 15°C did not differ between populations (desert: 6.39 ± 1.14 SE μmol m⁻³ s⁻¹, montane: 6.54 ± 1.13 SE μmol m⁻³ s⁻¹, P = 0.71) and declined with increasing stem diameter (P = 0.001). The temperature coefficient of respiration (Q ₁₀) varied seasonally within both environments (P = 0.05). Construction cost of stem sapwood was the same in both environments (desert: 1.46 ± 0.009 SE g glucose g⁻¹ sapwood, montane: 1.48 ± 0.009 SE glucose g⁻¹ sapwood, P = 0.14). Annual construction respiration calculated from construction cost, percent carbon and relative growth rate was greater in montane populations due to higher growth rates. These data provide no evidence of respiratory acclimation by desert trees. Estimated yearly stem maintenance respiration was greater in large desert trees than in large montane trees because of higher temperatures in the desert and because of increased allocation of biomass to sapwood. By analogy, these data suggest that under predicted increases in temperature and aridity, potential increases in aboveground carbon gain due to enhanced photosynthetic rates may be partially offset by increases in maintenance respiration in large trees growing in CO₂-enriched atmospheres. Received: 4 November 1996 / Accepted: 23 January 1997     

Annual soil respiration in broadleaf forests of northern Wisconsin: influence of moisture and site biological,chemical, and physical characteristics

Soil temperature and moisture influence soil respiration at a range of temporal and spatial scales. Although soil temperature and moisture may be seasonally correlated, intra and inter-annual variations in soil moisture do occur. There are few direct observations of the influence of local variation in species composition or other stand/site characteristics on seasonal and annual variations in soil moisture, and on cumulative annual soil carbon release. Soil climate and soil respiration from twelve sites in five different forest types were monitored over a 2-year period (1998–1999). Also measured were stand age, species composition, basal area, litter inputs, total above-ground wood production, leaf area index, forest floor mass, coarse and fine root mass, forest floor carbon and nitrogen concentration, root carbon and nitrogen concentration, soil carbon and nitrogen concentration, coarse fraction mass and volume, and soil texture. General soil respiration models were developed using soil temperature, daily soil moisture, and various site/soil characteristics. Of the site/soil characteristics, above-ground production, soil texture, roots + forest floor mass, roots + forest floor carbon:nitrogen, and soil carbon:nitrogen were significant predictors of soil respiration when used alone in respiration models; all of these site variables were weakly to moderately correlated with mean site soil moisture. Daily soil climate data were used to estimate the annual release of carbon (C) from soil respiration for the period 1998–1999. Mean annual soil temperature did not differ between the 2 years but mean annual soil moisture was approximately 9% lower in 1998 due to a summer drought. Soil C respired during 1998 ranged from 8.57 to 11.43 Mg C ha⁻¹ yr⁻¹ while the same sites released 10.13 and 13.57 Mg C ha⁻¹ yr⁻¹ in 1999; inter-annual differences of 15.41 and 15.73%, respectively. Among the 12 sites studied, we calculated that the depression of soil respiration linked to the drought caused annual differences of soil respiration from 11.00 to 15.78%. Annual estimates of respired soil C decreased with increasing site mean soil moisture. Similarly, the difference of respired carbon between the drought and the non-drought years generally decreased with increasing site mean soil moisture.     

Nitrogen Controls on Fine Root Substrate Quality in Temperate Forest Ecosystems

Nitrogen controls on fine root substrate quality (that is, nitrogen and carbon-fraction concentrations) were assessed using nitrogen availability gradients in the Harvard Forest chronic nitrogen addition plots, University of Wisconsin Arboretum, Blackhawk Island, Wisconsin, and New England spruce-fir transect. The 27 study sites encompassed within these four areas collectively represented a wide range of nitrogen availability (both quantity and form), soil types, species composition, aboveground net primary production, and climatic regimes. Changes in fine root substrate quality among sites were most frequently and strongly correlated with nitrate availability. For the combined data set, fine root nitrogen concentration increased (adjusted R ² = 0.46, P < 0.0001) with increasing site nitrate availability. Fine root “extractive” carbon-fraction concentrations decreased (adjusted R ² = 0.32, P < 0.0002), “acid-soluble” compounds increased (adjusted R ² = 0.35, P < 0.0001), and the “acid-insoluble” carbon fraction remained relatively high and stable (combined mean of 48.7 ± 3.1% for all sites) with increasing nitrate availability. Consequently, the ratio of acid-insoluble C–total N decreased (adjusted R ² = 0.40, P < 0.0001) along gradients of increasing nitrate availability. The coefficients of determination for significant linear regressions between site nitrate availability and fine root nitrogen and carbon-fraction concentrations were generally higher for sites within each of the four study areas. Within individual study sites, tissue substrate quality varied between roots in different soil horizons and between roots of different size classes. However, the temporal variation of fine root substrate quality indices within specific horizons was relatively low. The results of this study indicate that fine root substrate quality increases with increasing nitrogen availability and thus supports the substrate quality component of a hypothesized conceptual model of nitrogen controls on fine root dynamics that maintains that fine root production, mortality, substrate quality, and decomposition increase with nitrogen availability in forest ecosystems in a manner that is analogous to foliage.     

Supply-side controls on soil respiration among Oregon forests

To test the hypothesis that variation in soil respiration is related to plant production across a diverse forested landscape, we compared annual soil respiration rates with net primary production and the subsequent allocation of carbon to various ecosystem pools, including leaves, fine roots, forests floor, and mineral soil for 36 independent plots arranged as three replicates of four age classes in three climatically distinct forest types. Across all plots, annual soil respiration was not correlated with aboveground net primary production (R²=0.06, P>0.1) but it was moderately correlated with belowground net primary production (R²=0.46, P<0.001). Despite the wide range in temperature and precipitation regimes experienced by these forests, all exhibited similar soil respiration per unit live fine root biomass, with about 5 g of carbon respired each year per 1 g of fine root carbon (R²=0.45, P<0.001). Annual soil respiration was only weakly correlated with dead carbon pools such as forest floor and mineral soil carbon (R²=0.14 and 0.12, respectively). Trends between soil respiration, production, and root mass among age classes within forest type were inconsistent and do not always reflect cross‐site trends. These results are consistent with a growing appreciation that soil respiration is strongly influenced by the supply of carbohydrates to roots and the rhizosphere, and that some regional patterns of soil respiration may depend more on belowground carbon allocation than the abiotic constraints imposed on subsequent metabolism.