Takayuki Nakatsubo

Seasonal changes in the contribution of root respiration to total soil respiration in a cool-temperate deciduous forest

A trenching method was used to determine the contribution of root respiration to soil respiration. Soil respiration rates in a trenched plot (Rtrench) and in a control plot (Rcontrol) were measured from May 2000 to September 2001 by using an open-flow gas exchange system with an infrared gas analyser. The decomposition rate of dead roots (RD) was estimated by using a root-bag method to correct the soil respiration measured from the trenched plots for the additional decaying root biomass. The soil respiration rates in the control plot increased from May (240–320 mg CO2 m−2 h−1) to August (840–1150 mg CO2 m−2 h−1) and then decreased during autumn (200–650 mg CO2 m−2 h−1). The soil respiration rates in the trenched plot showed a similar pattern of seasonal change, but the rates were lower than in the control plot except during the 2 months following the trenching. Root respiration rate (Rr) and heterotrophic respiration rate (Rh) were estimated from Rcontrol, Rtrench, and RD. We estimated that the contribution of Rr to total soil respiration in the growing season ranged from 27 to 71%. There was a significant relationship between Rh and soil temperature, whereas Rr had no significant correlation with soil temperature. The results suggest that the factors controlling the seasonal change of respiration differ between the two components of soil respiration, Rr and Rh.

Effects of rainfall events on soil CO2 flux in a cool temperate deciduous broad-leaved forest

The effects of rainfall events on soil CO2 fluxes were examined in a cool temperate Quercus/Betula forest in Japan. The soil CO2 fluxes were measured using an open-flow gas exchange system with an infrared gas analyzer in the snow-free season from August 1999 to November 2000. Soil CO2 flux showed no significant diurnal trend on days without rain. In contrast, rainfall events caused a significant increase in soil CO2 flux. To determine the effect of rainfall events and to evaluate more precisely the daily and annual soil carbon flux, we constructed a multiple polynomial regression model that included two variables, soil temperature and soil water content, using the soil CO2 flux data recorded on sunny days. Daily soil carbon fluxes on sunny days calculated by the model were almost the same as those determined by the field measurements. On the contrary, the fluxes measured on rainy days were significantly higher than those calculated daily from the soil carbon fluxes by the model. Annual soil carbon fluxes in 1999 and 2000 were estimated using models that both do and do not take rainfall effects into consideration. The result indicates that post-rainfall increases in soil CO2 flux represent approximately 16–21% of the annual soil carbon flux in this cool temperate deciduous forest.

Contribution of micro-organisms to the carbon dynamics in black spruce (Picea mariana) forest soil in Canada

In order to clarify the role of micro-organisms in the carbon cycle of the boreal forest ecosystem, the vertical distribution of soil carbon, soil microbial biomass and respiratory activity was studied in a black spruce forest near Candle Lake in Saskatchewan, Canada. The total amount of carbon contained in moss and soil layers (to the depth of 50 cm beneath the mineral soil surface) was 7.2 kg m−2, about 47% of which was in the L and FH horizons of the soil. Soil microbial biomass per dry weight of soil was largest in the L horizon, while the biomass per ground area was largest in the FH horizon. Soil respiration rate, measured using a portable infrared gas analyzer, was highest in the FH horizon, exceeding 50% of the total soil respiration. Low but significant CO2 emission was detected even in deeper soil horizon (E horizon). We also examined the respiration rate of cut roots and the effect of root excision on respiration. The contribution of root respiration to total soil respiration, calculated from root biomass and respiration rate of cut roots, was about 54%. The amount of carbon evolved through microbial respiration during the snow-free season (June–October) was estimated as 221 g C m−2. Micro-organisms in the L horizon showed high respiratory activity as compared with those in deeper soil horizons.

Carbon and nitrogen limitation of soil microbial respiration in a High Arctic successional glacier foreland near Ny-Ålesund, Svalbard

The hypotheses that carbon and nitrogen availability limit microbial activity, and that the key factors limiting microbes vary along the successional gradient were tested in a High Arctic glacier foreland. We examined the responses of the respiration rate and the phospholipid fatty acid content to the addition of carbon and/or nitrogen. Soil samples were collected from the early stage and late stage of primary succession in the foreland of a glacier near Ny-Ålesund, Svalbard. The addition of both carbon (glucose) and nitrogen (ammonium nitrate) engendered an increase in the microbial respiration rate in the early stage of succession. In contrast, the addition of either carbon or nitrogen did not increase the microbial respiration rate. In the late stage of succession the addition of carbon alone, as well as the addition of both carbon and nitrogen, increased the microbial respiration rate. However, neither the addition of carbon nor the addition of nitrogen affected the total phospholipid fatty acid content (an index of microbial biomass) for any soil within 15 days of incubation at 10 ° C. An increase in the respiration rate was therefore attributed to changes in the physiological activities of the microbial community, such as enzymatic activity. Our study suggests that microbial respiration was limited by the low availability of both carbon and nitrogen in the early stage of succession. Thereafter, nitrogen limitation is mitigated.

Soil Microbial Biomass, Respiration Rate, and Temperature Dependence on a Successional Glacier Foreland in Ny-Alesund, Svalbard

Abstract We examined soil microbial activities, i.e., biomass, respiration rate, and temperature dependence of the respiration on a glacier foreland in Ny-Ålesund, Svalbard, Norway. We collected soil samples from 4 study sites that were set up along a primary succession (Site 1, the youngest, to Site 4, the oldest). Microbial biomass measured with the SIR method increased with successional age (55 to 724μg Cbiomass g−1 soil d.w. from Site 1 to Site 4). The microbial respiration rate of the soil was measured in a laboratory with an open-flow infrared gas-analyzer system, changing the temperature from 2° to 20°C at 3–4° intervals. The microbial respiration rate increased exponentially with the temperature at all sites. The temperature dependence (Q10) of the microbial respiration rate ranged from 2.2 to 4.1. The microbial respiration rates at a given temperature increased with succession as a step change (0.48, 0.43, 1.26, and 1.29μg C g−1soil h−1 at 8°C from Site 1 to Site 4, respectively). However, the substrate-specific respiration rate (respiration rate per gram soil carbon) decreased with successional age (0.034 to 0.006μg C mg−1Csoil h−1 from Site 1 to Site 4). A comparison of these respiratory properties with other ecosystems suggested that soil microorganisms in arctic soils have a high potential for decomposition when compared to those of other temperate ecosystems.

Production of biological soil crusts in the early stage of primary succession on a High Arctic glacier foreland

*We examined the photosynthetic characteristics and net primary production of biological soil crusts to evaluate their contribution to the carbon cycle in the High Arctic glacier foreland. *Biological soil crust samples were collected from a deglaciated area in Ny-Alesund, Svalbard, Norway. Net photosynthetic rates (Pn) and dark respiration rates (R) of biological soil crusts were determined using CO(2) gas exchange rates. We examined the effects of moisture conditions, temperature and photon flux density on Pn and R, and estimated the net primary production by a model based on the relationships between abiotic factors and Pn and R. *The maximum Pn value occurred at 50% of the maximum water-holding capacity. Pn decreased with increasing temperature and dropped below zero at high temperatures (c. > 13 degrees C). The estimated net primary production of the biological soil crust was greater than the net primary production of other vegetation when based on ground surface area, during the early stage of primary succession. Model simulation showed that the net primary production of the biological soil crust decreased with increasing temperature. *These results suggest that biological soil crust productivity plays an important role in the carbon cycle during the early stage of succession of the High Arctic glacier foreland, and is susceptible to temperature increases from global warming.

Comparative study of the mass loss rate of moss litter in boreal and subalpine forests in relation to temperature

The mossHylocomium splendens shows a very wide distribution in the Northern Hemisphere and may be useful as an indicator of climatic change on a global scale. We aimed to establish a convenient method to estimate the annual rate of litter mass loss of this species. The rate was calculated from the annual litter production rate and the amount of litter accumulated in the field. The litter production rate was estimated by analysis of the moss shoot growth. The rates calculated by this method tended to be larger than estimates obtained by the litter bag method. Using this method, we examined the difference in the litter mass loss rate along the altitudinal and latitudinal temperature gradients. The moss samples were collected from three boreal forests in Canada and four subalpine forests in Japan. At the subalpine sites, the annual rate of litter mass loss was within the range of 10–24% and tended to be smaller with increasing altitude. The rates in the boreal sites were similar to those in the subalpine sites despite lower mean annual temperatures. A significant log-linear relationship was observed between the annual mass loss rate and the cumulative value of monthly mean air temperatures higher than 0°C (CMT). Nitrogen concentration of the litter was positively correlated with mean annual air temperature. Site to site variation in the annual mass loss rate was largely explained by CMT and nitrogen concentration of the litter.

Successional changes in ectomycorrhizal fungi associated with the polar willow Salix polaris in a deglaciated area in the High Arctic, Svalbard

Polar willow (Salix polaris Wahlenb.), a mycorrhizal dwarf shrub, colonizes recently deglaciated areas in the High Arctic, Svalbard. To clarify successional changes in ECM fungi associated with S. polaris after glacier retreat, we examined the diversity and density of ECM fungi in culture and field conditions. Plant and soil samples were collected from three sites of different successional stages in the deglaciated area of Austre Brøggerbreen, near Ny-Ålesund, Svalbard. The successional stages were early stage with newly exposed bare ground (site I), transient stage with scattered colonization of Salix (sites IIa and IIb), and late stage with well-developed vegetation (site III). No ECM colonization on Salix was observed in soils collected from bare ground in early and transient stages (sites I and IIa). However, most Salix individuals showed ECM colonization in soils collected from sites close to Salix colonies in transient and late stages (sites IIb and III). Based on molecular analyses and operational taxonomic unit (OTU: >95% ITS sequence similarity) delimitations, we identified 15 OTUs/species in eight genera. The dominant OTU/species of ECM fungi identified in the transient and late stages was Geopora sp.1 and Cenococcum sp.1, respectively. In the culture experiment, ECM diversity was greater in late stage (eight OTUs/species) than in transient stage (three OTUs/species). This pattern was consistent with field observations, i.e., late-stage sites contained more OTUs/species of ECM fungi. These results indicate that species diversity of ECM fungi increases and the dominant species changes with the progress of succession after glacier retreat in the High Arctic.

Ecosystem development and carbon cycle on a glacier foreland in the high Arctic, Ny-Alesund, Svalbard

The Arctic terrestrial ecosystem is thought to be extremely susceptible to climate change. However, because of the diverse responses of ecosystem components to change, an overall response of the ecosystem carbon cycle to climate change is still hard to predict. In this review, we focus on several recent studies conducted to clarify the pattern of the carbon cycle on the deglaciated area of Ny-Ålesund, Svalbard in the high Arctic. Vegetation cover and soil carbon pools tended to increase with the progress of succession. However, even in the latter stages of succession, the size of the soil carbon pool was much smaller than those reported for the low Arctic tundra. Cryptogams contributed the major proportion of phytomass in the later stages. However, because of water limitation, their net primary production was smaller than that of the vascular plants. The compartment model that incorporated major carbon pools and flows suggested that the ecosystem of the later stages is likely to be a net sink of carbon at least for the summer season. Based on the eco-physiological characteristics of the major ecosystem components, we suggest several possible scenarios of future changes in the ecosystem carbon cycle.

Estimation of the biomass of fine roots and mycorrhizal fungi: a case study in a Japanese red pine (Pinus densiflora) stand

As part of a study on soil carbon flow in forest ecosystems, the biomass of fine roots (≪2.0 mm in diameter) and root-associated fungi, including ectomycorrhizal fungi, were estimated in the summer season in 1998 at a Pinus densiflora (Japanese red pine) stand in western Japan. Fine roots of pine were classified into three categories: class I roots (0.5–2.0 mm in diameter), long class II roots (long roots with diameter ≪0.5 mm; IIL), and short class II roots (short roots with diameter ≪0.5 mm; IIS). Total biomass of fine roots (I + IIL + IIS) at this stand was estimated to be 91.0 g m−2, about 23% of which was class II roots (IIL + IIS). Ergosterol, which is a component of fungal membranes, was analyzed to estimate the biomass of root-associated fungi in roots. In the upper soil layers (from the surface to 13.4 cm in depth), ergosterol contents in the class I, IIL and IIS roots were in the ranges 43.1–82.2, 126.1–196.3 and 271.2–321.0 µg g−1 root DW, respectively. The ergosterol content was converted to fungal biomass using the median (minimum–maximum) value of ergosterol concentration reported for ectomycorrhizal fungi. Root-associated fungal biomass in this stand was estimated to be 2.0 (0.5–9.6) g m−2. The data suggest the biomass of ectomycorrhizal fungi in the P. densiflora stand is small compared with that in other forest ecosystems.