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.     

Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil

The release of carbon as CO₂ from belowground processes accounts for about 70% of total ecosystem respiration. Insights about factors controlling soil CO₂ efflux are constrained by the challenge of apportioning sources of CO₂ between autotrophic tree roots (and mycorrhizal fungi) and heterotrophic microorganisms. In some temperate conifer forests, the reduction in soil CO₂ efflux after girdling (phloem removal) has been used to separate these sources. Girdling stops the flow of carbohydrates to the belowground portion of the ecosystem, which should slow respiration by roots and mycorrhizae while heterotrophic respiration should remain constant or be enhanced by the decomposition of newly dead roots. Therefore, the reduction in CO₂ efflux after girdling should be a conservative estimate of the belowground flux of C from trees. We tested this approach in two tropical Eucalyptus plantations. Tree canopies remained intact for more than 3 months after girdling, showing no reduction in light interception. The reduction in soil CO₂ efflux averaged 16–24% for the 3-month period after girdling. The reduction in CO₂ efflux was similar for plots with one half of the trees girdled and those with all of the trees girdled. Girdling did not reduce live fine root biomass for at least 5 months after treatment, indicating that large reserves of carbohydrates in the root systems of Eucalyptus trees maintained the roots and root respiration. Our results suggest that the girdling approach is unlikely to provide useful insights into the contribution of tree roots and heterotrophs to soil CO₂ efflux in this type of forest ecosystem.     

Differential controls by climate and substrate over the heterotrophic and rhizospheric components of soil respiration

The partitioning of soil respiration rates into the component processes of rhizospheric respiration (because of live roots and those microorganisms that subsist on root exudations) and heterotrophic respiration (because of decomposer microorganisms that subsist on the oxidation of soil organic matter) is difficult to accomplish through experimental observation. In order to minimize disturbance to the soil and maximize preservation of the natural relationships among roots, rhizospheric microorganisms, and decomposers, we conducted a girdling experiment in a subalpine forest dominated by lodgepole pine trees. In two separate years, we girdled trees in small forest plots (5–7 m in diameter) and trenched around the plots to sever invading roots in order to experimentally stop the transport of photosynthate from needles to roots, and eliminate rhizospheric respiration. Soil respiration rates in plots with trees girdled over 1 year prior to measurement were higher than those in plots with trees girdled 2–3 months prior to measurement. These results suggest that any stimulation of respiration because of the experimental artifact of fine root death and addition of labile carbon to the pool of decomposer substrates is slow, and occurs beyond the first growing season after girdling. Compared with control plots with nongirdled trees, soil respiration rates in plots with girdled trees were reduced by 31–44% at the mid‐summer respiratory maximum. An extreme drought during one of the 2 years used for observations caused greater reductions in the heterotrophic component of soil respiration compared with the rhizospheric component. In control plots, we observed a pulse in K₂SO₄‐extractable carbon during the spring snowmelt period, which was absent in plots with girdled trees. In control plots, soil microbial biomass increased from spring to summer, coincident with a seasonal increase in the rhizospheric component of soil respiration. In plots with girdled trees, the seasonal increase in microbial biomass was lower than in control plots. These results suggest that the observed seasonal increase in rhizospheric respiration rate in control plots was because of an increase in rhizospheric microbial biomass following ‘soil priming’ by a spring‐time pulse in dissolved organic carbon. Winter‐time, beneath‐snow microbial biomass was relatively high in control plots. Soil sucrose concentrations were approximately eight times higher during winter than during spring or summer, possibly being derived from the mechanical damage of shallow roots that use sucrose as protection against low‐temperature extremes. The winter‐time sucrose pulse was not observed in plots with girdled trees. The results of this study demonstrate that (1) the rhizospheric component of soil respiration rate at this site is significant in magnitude, (2) the heterotrophic component of soil respiration rate is more susceptible to seasonal drought than the rhizospheric component, and (3) the trees in this ecosystem exert a major control over soil carbon dynamics by ‘priming’ the soil with sugar exudates during the late‐spring snowmelt period and releasing high concentrations of sucrose to the soil during winter.     

Variations of root and heterotrophic respiration along environmental gradients in China's forests

Aims Root and heterotrophic respiration may respond differently to environmental variability, but little evidence is available from large-scale observations. Here we aimed to examine variations of root and heterotrophic respiration across broad geographic, climatic, soil and biotic gradients.Methods We conducted a synthesis of 59 field measurements on root and heterotrophic respiration across China's forests.Important findings Root and heterotrophic respiration varied differently with forest types, of which evergreen broadleaf forest was significantly different from those in other forest types on heterotrophic respiration but without statistically significant differences on root respiration. The results also indicated that root and heterotrophic respiration exhibited similar trends along gradients of precipitation, soil organic carbon and satellite-indicated vegetation growth. However, they exhibited different relationships with temperature: root respiration exhibited bimodal patterns along the temperature gradient, while heterotrophic respiration increased monotonically with temperature. Moreover, they showed different relationships with MOD17 GPP, with increasing trend observed for root respiration whereas insignificant change for heterotrophic respiration. In addition, root and heterotrophic respiration exhibited different changes along the age sequence, with insignificant change for root respiration and decreasing trend for heterotrophic respiration. Overall, these results suggest that root and heterotrophic respiration may respond differently to environmental variability. Our findings could advance our understanding on the different environmental controls of root and heterotrophic respiration and also improve our ability to predict soil CO₂ flux under a changing environment.     

Circadian rhythm of root pressure in popular and its driving factors

Aims Our main purposes were to investigate root pressure and its circadian rhythm of excised roots in ‘84K’ popular (Populus alba × P. glandulosa) cultured in soil and solution, to explore the influencing factors and their relationships with root pressure systematically and to understand the generation and rhythm regulation of root pressure. Methods We investigated the root pressure of excised roots in ‘84K’ popular using the method of digital pressure transducer. The diurnal rhythm of excised roots was conducted through different experimental treatments including sampling in different time, defoliation and girdling, together with ambient condition like soil temperature, differential or consistant temperature during day and night. Then we discussed the effects of root respiration and hydraulic conductivity on root pressure by further using chemical inhibitor. Furthermore, diurnal variation of osmotic potential and ions content as well as soluble sugar content of exudation was determined in order to explore their relationships with root pressure rhythm. Important findings Root pressure of excised roots in popular had diurnal rhythm which was higher during daytime and lower overnight. It reached its peak value in the morning to noon and valley value at 20:00. Root pressure of excised roots sampled at different time and cultured in different medium had influence on the rhythm of root pressure to some degrees, but did not the general rhythm of high in daytime and low overnight. Defoliation, girdling and the inhibitors for root respiration or cytomembrane hydraulic conductivity could affect the maximum value of root pressure while have no significant influence on the daily rhythm. Defoliation, girdling and respiration inhibitor reduced the maximum value of root pressure, whereas the hydraulic conductivity inhibitor had little influence on root pressure. The maximum value of root pressure declined with the decrease in soil temperature which could change the rhythm of root pressure. The synchronous change in the maximum value of root pressure and root respiration rate with temperature indicated that root respiration contributed to the change of root pressure along with temperature. Osmotic potential of root exudation was higher during the daytime and lower at night. Diurnal variations of ions and soluble sugar content of exudation were consistant with that of osmotic potential. The peak of root pressure measured under the condition of differential temperature during day and night was significant higher than that measured under constant temperature. In conclusion, root pressure of the poplar ‘84K’ showed significant diurnal rhythm, i.e. higher during the daytime and lower at night. The maximum value of root pressure was mainly regulated by root respiration metabolism. The factors such as respiration inhibitor, respiration substrate and temperature influence the value of the maximum root pressure of poplar ‘84K’. Root hydraulic conductivity had no significant influence on root pressure.     

Autotrophic and heterotrophic soil respiration in a Norway spruce forest: estimating the root decomposition and soil moisture effects in a trenching experiment

The two components of soil respiration, autotrophic respiration (from roots, mycorrhizal hyphae and associated microbes) and heterotrophic respiration (from decomposers), was separated in a root trenching experiment in a Norway spruce forest. In June 2003, cylinders (29.7 cm diameter) were inserted to 50 cm soil depth and respiration was measured both outside (control) and inside the trenched areas. The potential problems associated with the trenching treatment, increased decomposition of roots and ectomycorrhizal mycelia and changed soil moisture conditions, were handled by empirical modelling. The model was calibrated with respiration, moisture and temperature data of 2004 from the trenched plots as a training set. We estimate that over the first 5 months after the trenching, 45% of respiration from the trenched plots was an artefact of the treatment. Of this, 29% was a water difference effect and 16% resulted from root and mycelia decomposition. Autotrophic and heterotrophic respiration contributed to about 50% each of total soil respiration in the control plots averaged over the two growing seasons. We show that the potential problems with the trenching, decomposing roots and mycelia and soil moisture effects, can be handled by a modelling approach, which is an alternative to the sequential root harvesting technique.     

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.     

Seasonal fluctuations in Vitis vinifera root respiration in the field

We studied the seasonal fluctuation of soil respiration (R(S)), and its root-dependent (R(R)) and basal (R(B)) components, in a Vitis vinifera (Chardonnay) vineyard. The R(S) components were estimated through independent field methods (y-intercept and trenching) and modeled on the basis of a Q(10) response to soil temperature, and fine and coarse root respiration coefficients. The effect of assimilate availability on R(R) was assessed through a trunk girdling treatment. The apparent Q(10) for R(R) was twice that of R(B) (3.5 vs 1.6) and increased linearly with increasing vine root biomass. The fastest R(R) of fine roots was during rapid fruit growth and the fastest R(R) of coarse roots was immediately following fruit development. R(S) was estimated at 32.6 kg ha(-1) d(-1) (69% as a result of R(R) ) for the hottest month and at 7.6 kg ha(-1) d(-1) (18% as a result of R(R)) during winter dormancy. Annual R(S) was low compared with other natural and cultivated ecosystems: 5.4 Mg ha(-1) (46% as a result of R(R)). Our estimates of annual vineyard R(S) are the first for any horticultural crop and suggest that the assumption that they are similar to those of annual crops or forest trees might lead to an overestimation.     

Clean rain promotes fine root growth and soil respiration in a Norway spruce forest

Soil acidification and N saturation are considered to affect the decomposition of soil organic matter as well as growth and mortality of fine roots in many forest soils. Here we report from a field experiment where ‘clean rain’ has been applied to the soil for about 10 years under a roofed plot of a 71‐year‐old Norway spruce plantation at Solling, Central Germany. Reduced amounts of protons (?78%), sulphate (?53%), ammonium (?86%), and nitrate (?49%) were sprayed on the soil surface of the clean rain plot between 1992 and 2001. In an adjacent roofed control plot, throughfall was collected and immediately re‐sprinkled below the roof construction without any chemical manipulation. One year before the clean rain treatment started, live and dead fine root masses (≤2 mm) were determined from undisturbed soil cores down to 40 cm mineral soil depth. Total live fine root mass was significantly lower in the clean rain plot than in the control plot. After the first sampling, the soil holes were refilled with quartz sand and repeatedly sampled in June 1992, June 1996, and October 2001. There were no differences in live and dead fine root masses between the plots in 1992 and 1996. In 2001, both live and dead fine root masses of the clean rain plot were about twice as high as in the control plot, indicating that fine root growth recovered in the mineral soil following 10 years of clean rain treatment. Moreover, the clean rain treatment significantly reduced the total N concentrations of live fine roots and 1‐year‐old needles. Our results suggest that the reduced N input promoted fine root growth to compensate N deficiency. Reduced Al concentration in soil solution may have contributed to the recovery of fine root growth, however, the toxicity of Al species is largely unknown. Mean annual soil respiration rate was 24% higher in the period from 2000 to 2001, indicating that the clean rain treatment increased respiration of roots and heterotrophic microorganisms within the rhizosphere. Laboratory incubation of samples from the organic horizon and the top mineral soil revealed no differences between the plots in the decay rate of soil organic matter. Our results suggest that strong reductions in atmospheric N deposition from about 30 to 10 kg N ha^?1 yr^?1 and decreasing acid stress can have beneficial effects on growth of fine roots in the mineral soil within a decade. We conclude that biological recovery under reduced atmospheric loads can affect the nutrient and carbon budget of spruce soils in the long run.     

Effect of elevated atmospheric CO2 concentration on soil and root respiration in winter wheat by using a respiration partitioning chamber

Soil respiration in a cropland is the sum of heterotrophic (mainly microorganisms) and autotrophic (root) respiration. The contribution of both these types to soil respiration needs to be understood to evaluate the effects of environmental change on soil carbon cycling and sequestration. In this paper, the effects of free-air CO₂ enrichment (FACE) on hetero- and autotrophic respiration in a wheat field were differentiated and evaluated by a novel split-root growth and gas collection system. Elevated atmospheric pCO₂ of approximately 200 μmol mol⁻¹ above the ambient pCO₂ significantly increased soil respiration by 15.1 and 14.8% at high nitrogen (HN) and low nitrogen (LN) application rates, respectively. The effect of elevated atmospheric pCO₂ on root respiration was not consistent across the wheat growth stages. Elevated pCO₂ significantly increased and decreased root respiration at the booting-heading stage (middle stage) and the late-filling stage (late stage), respectively, in HN and LN treatments; however, no significant effect was found at the jointing stage (early stage). Thus, the effect of increased pCO₂ on cumulative root respiration for the entire wheat growing season was not significant. Cumulative root respiration accounted for approximately 25–30% of cumulative soil respiration in the entire wheat growing season. Consequently, cumulative microbial respiration (soil respiration minus root respiration) increased by 22.5 and 21.1% due to elevated pCO₂ in HN and LN, respectively. High nitrogen application significantly increased root respiration at the late stage under both elevated pCO₂ and ambient pCO₂; however, no significant effects were found on cumulative soil respiration, root respiration, and microbial respiration. These findings suggest that heterotrophic respiration, which is influenced by increased substrate supplies from the plant to the soil, is the key process to determine C emission from agro-ecosystems with regard to future scenarios of enriched pCO₂.     

The autotrophic contribution to soil respiration in a northern temperate deciduous forest and its response to stand disturbance

The goal of this study was to evaluate the contribution of oak trees (Quercus spp.) and their associated mycorrhizal fungi to total community soil respiration in a deciduous forest (Black Rock Forest) and to explore the partitioning of autotrophic and heterotrophic respiration. Trees on twelve 75 × 75-m plots were girdled according to four treatments: girdling all the oaks on the plot (OG), girdling half of the oak trees on a plot (O50), girdling all non-oaks on a plot (NO), and a control (C). In addition, one circular plot (diameter 50 m) was created where all trees were girdled (ALL). Soil respiration was measured before and after tree girdling. A conservative estimate of the total autotrophic contribution is approximately 50%, as indicated by results on the ALL and OG plots. Rapid declines in carbon dioxide (CO₂) flux from both the ALL and OG plots, 37 and 33%, respectively, were observed within 2 weeks following the treatment, demonstrating a fast turnover of recently fixed carbon. Responses from the NO and O50 treatments were statistically similar to the control. A non-proportional decline in respiration rates along the gradient of change in live aboveground biomass complicated partitioning of the overall rate of soil respiration and indicates that belowground carbon flux is not linearly related to aboveground disturbance. Our findings suggest that in this system there is a threshold disturbance level between 35 and 74% of live aboveground biomass loss, beyond which belowground dynamics change dramatically.     

Growth and maintenance respiration of roots of clonal Eucalyptus cuttings: scaling to stand-level

Root respiration consumes an important part of the daily assimilated carbon but the magnitude of this component of forest net ecosystem exchange and its partitioning among the different energy demanding processes in roots are still poorly documented. 5-month old Eucalyptus cuttings were grown in a greenhouse in pot filled with coarse sand. They were fertilized with three different amounts of a slow-release fertilizer with the doses of 8, 24 and 48 g of nitrogen per plant. Root respiration was measured using an infrared gas analyser by perfusing air through the pot on 9 plants per treatment on three dates 14 days apart. Measure of root respiration of the three treatments over time was made in order to obtain a large range of growth and nutrient uptake. Root respiration normalized at 22°C ranged from 0.09 to 0.23 g_C d^?1 for the three treatments during all the experiment. It was well predicted with a model that includes root growth rate and root nitrogen content.The nitrogen related maintenance coefficient was negatively correlated to the root nitrogen concentration suggesting a decrease in protein turnover with increasing fertility. Growth rate of fine root in a virtual stand was simulated using age-related allometric equations and further used to estimate root respiration in the field. Simulated root respiration increased over time from 0.39 to 3.14 g_C m^?2 d^?1 between 6 and 126 months assuming a turnover of 2 yr^?1 for fine roots. The major fraction of simulated root respiration in the field (78–92%) was used for the maintenance of the existing biomass.     

Ephemeral root modules in Fraxinus mandshurica

Historically, ephemeral roots have been equated with 'fine roots' (i.e. all roots of less than an arbitrary diameter, such as 2 mm), but evidence shows that 'fine roots' in woody species are complex branching systems with both rapid-cycling and slow-cycling components. A precise definition of ephemeral roots is therefore needed. Using a branch-order classification, a rhizotron method and sequential sampling of a root cohort, we tested the hypothesis that ephemeral root modules exist within the branching Fraxinus mandshurica (Manchurian ash) root system as distal nonwoody lateral branches, which show anatomical, nutritional and physiological patterns distinct from their woody mother roots. Our results showed that in F. mandshurica, distal nonwoody root branch orders die rapidly as intact lateral branches (or modules). These nonwoody branch orders exhibited highly synchronous changes in tissue nitrogen concentrations and respiration, dominated root turnover, nutrient flux and root respiration, and never underwent secondary development. The ephemeral root modules proposed here may provide a functional basis for differentiating and sampling short-lived absorptive roots in woody plants, and represent a conceptual leap over the traditional coarse-fine root dichotomies based on arbitrary size classes.     

Relationships between fine root dynamics and nitrogen availability in Michigan northern hardwood forests

Minirhizotrons were used to observe fine root (Б mm) production, mortality, and longevity over 2 years in four sugar-maple-dominated northern hardwood forests located along a latitudinal temperature gradient. The sites also differed in N availability, allowing us to assess the relative influences of soil temperature and N availability in controlling fine root lifespans. Root production and mortality occurred throughout the year, with most production occurring in the early portion of the growing season (by mid-July). Mortality was distributed much more evenly throughout the year. For surface fine roots (0-10 cm deep), significant differences in root longevity existed among the sites, with median root lifespans for root cohorts produced in 1994 ranging from 405 to 540 days. Estimates of fine root turnover, based on the average of annual root production and mortality as a proportion of standing crop, ranged from 0.50 to 0.68 year^-1 for roots in the upper 30 cm of soil. The patterns across sites in root longevity and turnover did not follow the north to south temperature gradient, but rather corresponded to site differences in N availability, with longer average root lifespans and lower root turnover occurring where N availability was greater. This suggests the possibility that roots are maintained as long as the benefit (nutrients) they provide outweighs the C cost of keeping them alive. Root N concentrations and respiration rates (at a given temperature) were also higher at sites where N availability was greater. It is proposed that greater metabolic activity for roots in nitrogen-rich zones leads to greater carbohydrate allocation to those roots, and that a reduction in root C sink strength when local nutrients are depleted provides a mechanism through which root lifespan is regulated in these forests.     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

The effect of phosphorus availability on the carbon economy of contrasting common bean (Phaseolus vulgaris L.) genotypes

A common response to low phosphorus availability is increased relative biomass allocation to roots. The resulting increase in root:shoot ratio presumably enhances phosphorus acquisition, but may also reduce growth rates by diverting carbon to the production of heterotrophic rather than photosynthetic tissues. To assess the importance of increased carbon allocation to roots for the adaptation of plants to low P availability, carbon budgets were constructed for four common bean genotypes with contrasting adaptation to low phosphorus availability in the field ("phosphorus efficiency"). Solid-phase-buffered silica sand provided low (1 microM), medium (10 microM), and high (30 microM) phosphorus availability. Compared to the high phosphorus treatment, plant growth was reduced by 20% by medium phosphorus availability and by more than 90% by low phosphorus availability. Low phosphorus plants utilized a significantly larger fraction of their daytime net carbon assimilation on root respiration (c. 40%) compared to medium and high phosphorus plants (c. 20%). No significant difference was found among genotypes in this respect. Genotypes also had similar rates of P absorption per unit root weight and plant growth per unit of P absorbed. However, P-efficient genotypes allocated a larger fraction of their biomass to root growth, especially under low P conditions. Efficient genotypes had lower rates of root respiration than inefficient genotypes, which enabled them to maintain greater root biomass allocation than inefficient genotypes without increasing overall root carbon costs.     

Soil plus root respiration and microbial biomass following water, nitrogen, and phosphorus application at a high arctic semi desert

In order to investigate the effects of anticipated increased precipitation and changing soil nutrient levels on soil CO₂ efflux from high arctic semi desert, a field experiment was carried out in Northeast Greenland. Water, phosphorus, and nitrogen were added to plots in a fully factorial design. Soil microbial biomass carbon was analysed after one year, and respiration from soil plus roots was measured in situ throughout the third growing season after initiation of the experiment. Soil plus root respiration was enhanced by up to 47%, and the microbial biomass by 24%, by the weekly water additions, but not by nutrient additions. The direct effect of increased soil moisture on CO₂ efflux suggests that future changes of precipitation levels and patterns may strongly affect below-ground respiration in arctic semi deserts, with direction of responses depending upon amounts and frequencies of precipitation events. Morover, low CO₂ emission at low light intensities regardless of treatment suggests that the major part of the below-ground respiration originated from turnover of recently fixed C. Hence, the more recalcitrant soil organic matter C pool may not change in proportion to changes in below-ground respiration rate.     

Partitioning of soil respiration components and evaluating the mycorrhizal contribution to soil respiration in a semiarid grassland

土壤呼吸组分的区分对于理解地下碳循环过程非常重要。而菌根真菌在地下碳循环过程中扮演着重要的角色, 但是有关菌根呼吸在草原生态系统中的研究相对较少。该研究在内蒙古半干旱草原应用深浅环网孔法, 结合浅环、深环(排除根系)和一个带有40 μm孔径窗口的土壤环(排除根系但是有菌根菌丝体)将根和菌丝物理分离, 来区分不同的呼吸组分。结果表明: 异养呼吸对总呼吸的贡献比例为51%, 根呼吸的贡献比例为26%, 菌根呼吸的贡献比例为23%, 菌根呼吸的比例3年变化范围为21%-26%。与国内外研究相比, 此方法提供了一个相对稳定的菌根呼吸测量精度范围, 在草原生态系统中切实可行。对菌根呼吸的准确定量将有助于预测草原生态系统土壤碳释放过程对未来气候变化的响应。     

半干旱草原土壤呼吸组分区分与菌根呼吸的贡献

土壤呼吸组分的区分对于理解地下碳循环过程非常重要。而菌根真菌在地下碳循环过程中扮演着重要的角色, 但是有关菌根呼吸在草原生态系统中的研究相对较少。该研究在内蒙古半干旱草原应用深浅环网孔法, 结合浅环、深环(排除根系)和一个带有40 μm孔径窗口的土壤环(排除根系但是有菌根菌丝体)将根和菌丝物理分离, 来区分不同的呼吸组分。结果表明: 异养呼吸对总呼吸的贡献比例为51%, 根呼吸的贡献比例为26%, 菌根呼吸的贡献比例为23%, 菌根呼吸的比例3年变化范围为21%-26%。与国内外研究相比, 此方法提供了一个相对稳定的菌根呼吸测量精度范围, 在草原生态系统中切实可行。对菌根呼吸的准确定量将有助于预测草原生态系统土壤碳释放过程对未来气候变化的响应。     

Soil carbon cycling in a temperate forest: radiocarbon-based estimates of residence times, sequestration rates and partitioning of fluxes

Temperate forests of North America are thought to besignificant sinks of atmospheric CO₂. Wedeveloped a below-ground carbon (C) budget forwell-drained soils in Harvard Forest Massachusetts, anecosystem that is storing C. Measurements of carbonand radiocarbon (¹⁴C) inventory were used todetermine the turnover time and maximum rate ofCO₂ production from heterotrophic respiration ofthree fractions of soil organic matter (SOM):recognizable litter fragments (L), humified lowdensity material (H), and high density ormineral-associated organic matter (M). Turnover timesin all fractions increased with soil depth and were2–5 years for recognizable leaf litter, 5–10 years forroot litter, 40–100+ years for low density humifiedmaterial and >100 years for carbon associated withminerals. These turnover times represent the timecarbon resides in the plant + soil system, and mayunderestimate actual decomposition rates if carbonresides for several years in living root, plant orwoody material.Soil respiration was partitioned into two componentsusing ¹⁴C: recent photosynthate which ismetabolized by roots and microorganisms within a yearof initial fixation (Recent-C), and C that is respiredduring microbial decomposition of SOM that resides inthe soil for several years or longer (Reservoir-C).For the whole soil, we calculate that decomposition ofReservoir-C contributes approximately 41% of thetotal annual soil respiration. Of this 41%,recognizable leaf or root detritus accounts for 80%of the flux, and 20% is from the more humifiedfractions that dominate the soil carbon stocks.Measurements of CO₂ and ¹⁴CO₂ in thesoil atmosphere and in total soil respiration werecombined with surface CO₂ fluxes and a soil gasdiffusion model to determine the flux and isotopicsignature of C produced as a function of soil depth. 63% of soil respiration takes place in the top 15 cmof the soil (O + A + Ap horizons). The average residencetime of Reservoir-C in the plant + soil system is8±1 years and the average age of carbon in totalsoil respiration (Recent-C + Reservoir-C) is 4±1years.The O and A horizons have accumulated 4.4 kgC m^–2above the plow layer since abandonment by settlers inthe late-1800's. C pools contributing the most to soilrespiration have short enough turnover times that theyare likely in steady state. However, most C is storedas humified organic matter within both the O and Ahorizons and has turnover times from 40 to 100+ yearsrespectively. These reservoirs continue to accumulatecarbon at a combined rate of 10–30 gC m^{minus 2}yr^–1. This rate of accumulation is only 5–15% of the total ecosystem C sink measured in this stand using eddy covariance methods.