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.     

Influence of interactions between litter decomposition and rhizosphere activity on soil respiration and on the temperature sensitivity in a subtropical montane forest in SW China

Aims

The aims were to identify the effects of interactions between litter decomposition and rhizosphere activity on soil respiration and on the temperature sensitivity of soil respiration in a subtropical forest in SW China.

Methods

Four treatments were established: control (CK), litter removal (NL), trenching (NR) and trenching together with litter removal (NRNL). Soil CO₂ efflux, soil temperature, and soil water content were measured once a month over two years. Soil respiration was divided into four components: the decomposition of basic soil organic matter (SOM), litter respiration, root respiration, and the interaction effect between litter decomposition and rhizosphere activity. A two-factor regression equation was used to correct the value of soil CO₂ efflux.

Results

We found a significant effect of the interaction between litter decomposition and rhizosphere activity (R _INT) on total soil respiration, and R _INT exhibited significant seasonal variation, accounting for 26 and 31 % of total soil respiration in the dry and rainy seasons, respectively. However, we found no significant interaction effect on the temperature sensitivity of soil respiration. The temperature sensitivity was significantly increased by trenching compared with the control, but was unchanged by litter removal.

Conclusions

Though the interaction between litter decomposition and rhizosphere activity had no effects on temperature sensitivity, it had a significant positive effect on soil respiration. Our results not only showed strong influence of rhizosphere activity on temperature sensitivity, but provided a viable way to identify the contribution of SOM to soil respiration, which could help researchers gain insights on the carbon cycle.     

Contribution of aboveground litter,belowground litter,and rhizosphere respiration to total soil CO<Subscript>2</Subscript> efflux in an old growth coniferous forest

In an old growth coniferous forest located in the central Cascade Mountains, Oregon, we added or removed aboveground litter and terminated live root activity by trenching to determine sources of soil respiration. Annual soil efflux from control plots ranged from 727 g C m⁻² year⁻¹ in 2002 to 841 g C m⁻² year⁻¹ in 2003. We used aboveground litter inputs (149.6 g C m⁻² year⁻¹) and differences in soil CO₂ effluxes among treatment plots to calculate contributions to total soil efflux by roots and associated rhizosphere organisms and by heterotrophic decomposition of organic matter derived from aboveground and belowground litter. On average, root and rhizospheric respiration (R_r) contributed 23%, aboveground litter decomposition contributed 19%, and belowground litter decomposition contributed 58% to total soil CO₂ efflux, respectively. These values fall within the range of values reported elsewhere, although our estimate of belowground litter contribution is higher than many published estimates, which we argue is a reflection of the high degree of mycorrhizal association and low nutrient status of this ecosystem. Additionally, we found that measured fluxes from plots with doubled needle litter led to an additional 186 g C m⁻² year⁻¹ beyond that expected based on the amount of additional carbon added; this represents a priming effect of 187%, or a 34% increase in the total carbon flux from the plots. This finding has strong implications for soil C storage, showing that it is inaccurate to assume that increases in net primary productivity will translate simply and directly into additional belowground storage.     

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.     

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.     

Effects of Litter Inputs on N<Subscript>2</Subscript>O Emissions from a Tropical Rainforest in Southwest China

Litter inputs are expected to have a strong impact on soil N₂O efflux. This study aimed to assess the effects of the litter decomposition process and nutrient efflux from litter to soil on soil N₂O efflux in a tropical rainforest. A paired study with a control (L) treatment and a litter-removed (NL) treatment was followed for 2 years, continuously monitoring the effects of these treatments on soil N₂O efflux, fresh litter input, decomposed litter carbon (LCI) and nitrogen (LNI), soil nitrate (NO₃ ^?–N), ammonium (NH₄ ⁺–N), dissolved organic carbon (DOC), and dissolved nitrogen (DN). Soil N₂O flux was 0.48 and 0.32 kg N₂O–N ha^?1 year^?1 for the L and NL treatments, respectively. Removing the litter caused a decrease in the annual soil N₂O emission by 33%. The flux values from the litter layer were higher in the rainy season as compared to the dry season (2.10 ± 0.28 vs. 1.44 ± 0.35 μg N m^?2 h^?1). The N₂O fluxes were significantly correlated with the soil NO₃ ^?–N contents (P < 0.05), indicating that the N₂O emission was derived mainly from denitrification as well as other NO₃ ^? reduction processes. Suitable soil temperature and moisture sustained by rainfall were jointly attributed to the higher soil N₂O fluxes of both treatments in the rainy season. The N₂O fluxes from the L were mainly regulated by LCI, whereas those from the NL were dominated jointly by soil NO₃ ^? content and temperature. The effects of LCI and LNI on the soil N₂O fluxes were the greatest in the 2 months after litter decomposition. Our results show that litter may affect not only the variability in the quantity of N₂O emitted, but also the mechanisms that govern N₂O production. However, further studies are still required to elucidate the impacting mechanisms of litter decomposition on N₂O emission from tropical forests.     

Partitioning sources of soil-respired CO2 and their seasonal variation using a unique radiocarbon tracer

Soil respiration is derived from heterotrophic (decomposition of soil organic matter) and autotrophic (root/rhizosphere respiration) sources, but there is considerable uncertainty about what factors control variations in their relative contributions in space and time. We took advantage of a unique whole‐ecosystem radiocarbon label in a temperate forest to partition soil respiration into three sources: (1) recently photosynthesized carbon (C), which dominates root and rhizosphere respiration; (2) leaf litter decomposition and (3) decomposition of root litter and soil organic matter >1–2 years old. Heterotrophic sources and specifically leaf litter decomposition were large contributors to total soil respiration during the growing season. Relative contributions from leaf litter decomposition ranged from a low of ～1±3% of total soil respiration (6± 3 mg C m^?2 h^?1) when leaf litter was extremely dry, to a high of 42±16% (96± 38 mg C m^?2 h^?1). Total soil respiration fluxes varied with the strength of the leaf litter decomposition source, indicating that moisture‐dependent changes in litter decomposition drive variability in total soil respiration fluxes. In the surface mineral soil layer, decomposition of C fixed in the original labeling event (3–5 years earlier) dominated the isotopic signature of heterotrophic respiration. Root/rhizosphere respiration accounted for 16±10% to 64±22% of total soil respiration, with highest relative contributions coinciding with low overall soil respiration fluxes. In contrast to leaf litter decomposition, root respiration fluxes did not exhibit marked temporal variation ranging from 34±14 to 40±16 mg C m^?2 h^?1 at different times in the growing season with a single exception (88±35 mg C m^?2 h^?1). Radiocarbon signatures of root respired CO₂ changed markedly between early and late spring (March vs. May), suggesting a switch from stored nonstructural carbohydrate sources to more recent photosynthetic products.     

Elevated CO2 and temperature impacts on different components of soil CO2 efflux in Douglas-fir terracosms

Although numerous studies indicate that increasing atmospheric CO₂ or temperature stimulate soil CO₂ efflux, few data are available on the responses of three major components of soil respiration [i.e. rhizosphere respiration (root and root exudates), litter decomposition, and oxidation of soil organic matter] to different CO₂ and temperature conditions. In this study, we applied a dual stable isotope approach to investigate the impact of elevated CO₂ and elevated temperature on these components of soil CO₂ efflux in Douglas-fir terracosms. We measured both soil CO₂ efflux rates and the ¹³C and ¹⁸O isotopic compositions of soil CO₂ efflux in 12 sun-lit and environmentally controlled terracosms with 4-year-old Douglas fir seedlings and reconstructed forest soils under two CO₂ concentrations (ambient and 200 ppmv above ambient) and two air temperature regimes (ambient and 4 °C above ambient). The stable isotope data were used to estimate the relative contributions of different components to the overall soil CO₂ efflux. In most cases, litter decomposition was the dominant component of soil CO₂ efflux in this system, followed by rhizosphere respiration and soil organic matter oxidation. Both elevated atmospheric CO₂ concentration and elevated temperature stimulated rhizosphere respiration and litter decomposition. The oxidation of soil organic matter was stimulated only by increasing temperature. Release of newly fixed carbon as root respiration was the most responsive to elevated CO₂, while soil organic matter decomposition was most responsive to increasing temperature. Although some assumptions associated with this new method need to be further validated, application of this dual-isotope approach can provide new insights into the responses of soil carbon dynamics in forest ecosystems to future climate changes.     

Components of forest soil CO2 efflux estimated from Δ14C values of soil organic matter

Aims

The partitioning of the total soil CO₂ efflux into its two main components: respiration from roots (and root-associated organisms) and microbial respiration (by means of soil organic matter (SOM) and litter decomposition), is a major need in soil carbon dynamics studies in order to understand if a soil is a net sink or source of carbon.

Methods

The heterotrophic component of the CO₂ efflux was estimated for 11 forest sites as the ratio between the carbon stocks of different SOM pools and previously published (Δ¹⁴C derived) turnover times. The autotrophic component, including root and root-associated respiration, was calculated by subtracting the heterotrophic component from total soil chamber measured CO₂ efflux.

Results

Results suggested that, on average, 50.4 % of total soil CO₂ efflux was derived from the respiration of the living roots, 42.4 % from decomposition of the litter layers and less than 10 % from decomposition of belowground SOM.

Conclusions

The Δ¹⁴C method proved to be an efficient tool by which to partition soil CO₂ efflux and quantify the contribution of the different components of soil respiration. However the average calculated heterotrophic respiration was statistically lower compared with two previous studies dealing with soil CO₂ efflux partitioning (one performed in the same study area; the other a meta-analysis of soil respiration partitioning). These differences were probably due to the heterogeneity of the SOM fraction and to a sub-optimal choice of the litter sampling period.     

Tropical tree species composition affects the oxidation of dissolved organic matter from litter

Plant species effects on soil nutrient availability are relatively well documented, but the effects of species differences in litter chemistry on soil carbon cycling are less well understood, especially in the species-rich tropics. In many wet tropical forest ecosystems, leaching of dissolved organic matter (DOM) from the litter layer accounts for a significant proportion of litter mass loss during decomposition. Here we investigated how tree species differences in soluble dissolved organic C (DOC) and nutrients affected soil CO₂ fluxes in laboratory incubations. We leached DOM from freshly fallen litter of six canopy tree species collected from a tropical rain forest in Costa Rica and measured C-mineralization. We found significant differences in litter solubility and nutrient availability. Following DOM additions to soil, rates of heterotrophic respiration varied by as much as an order of magnitude between species, and overall differences in total soil CO₂ efflux varied by more than four-fold. Variation in the carbon: phosphorus ratio accounted for 51% of the variation in total CO₂ flux between species. These results suggest that tropical tree species composition may influence soil C storage and mineralization via inter-specific variation in plant litter chemistry.     

Spatial and temporal variation in soil CO2 efflux in an old-growth neotropical rain forest, La Selva, Costa Rica

Our objectives were to quantify and compare soil CO₂ efflux of two dominant soil types in an old-growth neotropical rain forest in the Atlantic zone of Costa Rica, and to evaluate the control of environmental factors on CO₂ release. We measured soil CO₂ efflux from eight permanent soil chambers on six Oxisol sites. Three sites were developed on old river terraces (old alluvium) and the other three were developed on old lava flows (residual). At the same time we measured soil CO₂ concentrations, soil water content and soil temperature at various depths in 6 soil shafts (3 m deep). Between old alluvium sites, the two-year average CO₂ flux rates ranged from 117.3 to 128.9 mg C m^–2 h^–1. Significantly higher soil CO₂ flux occurred on the residual sites (141.1 to 184.2 mg C m^–2 h^–1). Spatial differences in CO₂ efflux were related to fine root biomass, soil carbon and phosphorus concentration but also to soil water content. Spatial variability in CO₂ storage was high and the amount of CO₂ stored in the upper and lower soil profile was different between old alluvial and residual sites. The major factor identified for explaining temporal variations in soil CO₂ efflux was soil water content. During periods of high soil water content CO₂ emission decreased, probably due to lower diffusion and CO₂ production rates. During the 2-year study period inter-annual variation in soil CO₂ efflux was not detected.     

Time-dependent responses of soil CO2 efflux components to elevated atmospheric [CO2] and temperature in experimental forest mesocosms

We previously used dual stable isotope techniques to partition soil CO₂ efflux into three source components (rhizosphere respiration, litter decomposition, and soil organic matter (SOM) oxidation) using experimental chambers planted with Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] seedlings. The components responded differently to elevated CO₂ (ambient + 200 mol mol^–1) and elevated temperature (ambient + 4 °C) treatments during the first year. Rhizosphere respiration increased most under elevated CO₂, and SOM oxidation increased most under elevated temperature. However, many studies show that plants and soil processes can respond to altered climates in a transient way. Herein, we extend our analysis to 2 years to evaluate the stability of the responses of the source components. Total soil CO₂ efflux increased significantly under elevated CO₂ and elevated temperature in both years (1994 and 1995), but the enhancement was much less in 1995. Rhizosphere respiration increased less under elevated temperature in 1995 compared with 1994. Litter decomposition also tended to increase comparatively less in 1995 under elevated CO₂, but was unresponsive to elevated temperature between years. In contrast, SOM oxidation was similar under elevated CO₂ in the 2 years. Less SOM oxidation occurred under elevated temperature in 1995 compared with 1994. Our results indicate that temporal variations can occur in CO₂ production by the sources. The variations likely involve responses to antecedent physical disruption of the soil and physiological processes.     

Increases in soil respiration following labile carbon additions linked to rapid shifts in soil microbial community composition

Organic matter decomposition and soil CO₂ efflux are both mediated by soil microorganisms, but the potential effects of temporal variations in microbial community composition are not considered in most analytical models of these two important processes. However, inconsistent relationships between rates of heterotrophic soil respiration and abiotic factors, including temperature and moisture, suggest that microbial community composition may be an important regulator of soil organic matter (SOM) decomposition and CO₂ efflux. We performed a short-term (12-h) laboratory incubation experiment using tropical rain forest soil amended with either water (as a control) or dissolved organic matter (DOM) leached from native plant litter, and analyzed the effects of the treatments on soil respiration and microbial community composition. The latter was determined by constructing clone libraries of small-subunit ribosomal RNA genes (SSU rRNA) extracted from the soil at the end of the incubation experiment. In contrast to the subtle effects of adding water alone, additions of DOM caused a rapid and large increase in soil CO₂ flux. DOM-stimulated CO₂ fluxes also coincided with profound shifts in the abundance of certain members of the soil microbial community. Our results suggest that natural DOM inputs may drive high rates of soil respiration by stimulating an opportunistic subset of the soil bacterial community, particularly members of the Gammaproteobacteria and Firmicutes groups. Our experiment indicates that variations in microbial community composition may influence SOM decomposition and soil respiration rates, and emphasizes the need for in situ studies of how natural variations in microbial community composition regulate soil biogeochemical processes.     

Seasonal changes and vertical distribution of root standing biomass of graminoids and shrubs at a Siberian tundra site

Background &; aims

Elevated atmospheric CO₂ (eCO₂) can affect soil-plant systems via stimulating plant growth, rhizosphere activity and the decomposition of added (crop residues) or existing (priming) soil organic carbon (C). Increases in C inputs via root exudation, rhizodeposition and root turnover are likely to alter the decomposition of crop residues but will ultimately depend on the N content of the residues and the soil.

Methods

Two soil column experiments were conducted under ambient CO₂ (aCO₂, 390 ppm) and eCO₂ (700 ppm) in a glasshouse using dual-labelled (¹³C/¹⁵N) residues of wheat (Triticum aestivum cv. Yitpi) and field pea (Pisum sativum L. cv. PBA Twilight). The effects of eCO₂ and soil N status on wheat rhizosphere activity and residue decomposition and also N recovery from crop residues with different N status (C/N ratio 19.4–115.4) by different plant treatments (wheat, wheat + 25 mg N kg^?1 and field pea).

Results

Total belowground CO₂ efflux was enhanced under eCO₂ despite no increases in root biomass. Plants decreased residue decomposition, indicating a negative rhizosphere effect. For wheat, eCO₂ reduced the negative rhizosphere effect, resulting in greater rates of decomposition and recovery of N from field pea residues, but only when N fertiliser was added. For field pea, eCO₂ enhanced the negative rhizosphere effect resulting in lower decomposition rates and N recovery from field pea residue.

Conclusions

The effect of eCO₂ on N utilisation varied with the type of residue, enhancing N utilisation of wheat but repressing that of field pea residues, which in turn could alter the amount of N supplied to subsequent crops. Furthermore, reduced decomposition of residues under eCO₂ may slow the formation of new soil C and have implications for long-term soil fertility.

     

Interactive effects of soil nitrogen and atmospheric carbon dioxide on root/rhizosphere carbon dioxide efflux from loblolly and ponderosa pine seedlings

We measured CO₂ efflux from intact root/rhizosphere systems of 155 day old loblolly (Pinus taeda L.) and ponderosa (Pinus ponderosa Dougl. ex Laws.) pine seedlings in order to study the effects of elevated atmospheric CO₂ on the below-ground carbon balance of coniferous tree seedlings. Seedlings were grown in sterilized sand culture, watered daily with either 1, 3.5 or 7 mt M NH ₄ ⁺ , and maintained in an atmosphere of either 35 or 70 Pa CO₂. Carbon dioxide efflux (mol CO₂ plant^–1 s^–1) from the root/rhizosphere system of both species significantly increased when seedlings were grown in elevated CO₂, primarily due to large increases in root mass. Specific CO₂ efflux (mol CO₂ g root^–1 s^–1) responded to CO₂ only under conditions of adequate soil nitrogen availability (3.5 mt M). Under these conditions, CO₂ efflux rates from loblolly pine increased 70% from 0.0089 to 0.0151 mol g^–1 s^–1 with elevated CO₂ while ponderosa pine responded with a 59% decrease, from 0.0187 to 0.0077 mol g^–1 s^–1. Although below ground CO₂ efflux from seedlings grown in either sub-optimal (1 mt M) or supra-optimal (7 mt M) nitrogen availability did not respond to CO₂, there was a significant nitrogen treatment effect. Seedlings grown in supra-optimal soil nitrogen had significantly increased specific CO₂ efflux rates, and significantly lower total biomass compared to either of the other two nitrogen treatments. These results indicate that carbon losses from the root/rhizosphere systems are responsive to environmental resource availability, that the magnitude and direction of these responses are species dependent, and may lead to significantly different effects on whole plant carbon balance of these two forest tree species.     

Photosynthesis controls of CO2 efflux from maize rhizosphere

The effects of different shading periods of maize plants on rhizosphere respiration and soil organic matter decomposition were investigated by using a ¹³C natural abundance and ¹⁴C pulse labeling simultaneously. ¹³C was a tracer for total C assimilated by maize during the whole growth period, and ¹⁴C was a tracer for recently assimilated C. CO₂ efflux from bare soil was 4 times less than the total CO₂ efflux from planted soil under normal lighting. Comparing to the normal lighting control (12/12 h day/night), eight days with reduced photosynthesis (12/36 h day/night period) and strongly reduced photosynthesis (12/84 h day/night period) resulted in 39% and 68% decrease of the total CO₂ efflux from soil, respectively. The analysis of ¹³C natural abundance showed that root-derived CO₂ efflux accounted for 82%, 68% and 56% of total CO₂ efflux from the planted soil with normal, prolonged and strongly prolonged night periods, respectively. Clear diurnal dynamics of the total CO₂ efflux from soil with normal day-night period as well as its strong reduction by prolonged night period indicated tight coupling with plant photosynthetic activity. The light-on events after prolonged dark periods led to increases of root-derived and therefore of total CO₂ efflux from soil. Any factor affecting photosynthesis, or substrate supply to roots and rhizosphere microorganisms, is an important determinant of root-derived CO₂ efflux, and thereby, total CO₂ efflux from soils. ¹⁴C labeling of plants before the first light treatment did not show any significant differences in the ¹⁴CO₂ respired in the rhizosphere between different dark periods because the assimilate level in the plants was high. Second labeling, conducted after prolonged night phases, showed higher contribution of recently assimilated C (¹⁴C) to the root-derived CO₂ efflux by shaded plants. Results from ¹³C natural abundance showed that the cultivation of maize on Chromic Luvisol decreased soil organic matter (SOM) mineralization compared to unplanted soil (negative priming effect). A more important finding is the observed tight coupling of the negative rhizosphere effect on SOM decomposition with photosynthesis.     

Changes in Photosynthesis and Fruit Characteristics in Olive in Response to Assimilate Availability

The source level in the olive cultivar Leccino was varied by girdling at different stages of fruit growth. Afterwards, the effects on gas exchange, fruit growth, and ripening and blooming were studied. Girdling during fruit growth did not significantly influence net photosynthetic rate (P _N) except in the last phase of fruit growth when the P _N was reduced. In the girdled branch, P _N began to decrease at the onset of starch accumulation because fruit growth ceased. In mid-November stomatal conductance (g _s) and transpiration rate (E) were also reduced by girdling, whereas sub-stomatal CO₂ concentration (C _i) increased in leaves from the girdled branches. The total chlorophyll content (Chl) tended to decrease in parallel with the reduced P _N. Girdling did not substantially influence the leaf and shoot water contents. The large availability of assimilates seems to cause an earlier fruit ripening. In general, girdling increased fruit dry mass. Healing before the time when the majority of pulp growth occurs reduced the effect of girdling. June girdling increased the pit dry mass. Girdling at the beginning of August and September, compared to the control, increased the pulp dry mass, but the pit dry mass did not differ with respect to the control. The percentage of oil in the fruit, on a dry mass basis, increased with August and September girdlings, but the percentage of oil in the pulp did not change. Girdling reduced shoot growth, but the internode length was unchanged. Girdling slightly stimulated differentiation of flower buds.     

Soil CO<Subscript><Emphasis Type="Bold">2</Emphasis></Subscript> efflux and fungal and bacterial biomass in a plantation and a secondary forest in wet tropics in Puerto Rico

We examined the effects of root and litter exclusion on the rate of soil CO₂ efflux and microbial biomass using trenching and tent separation techniques in a secondary forest (SF) and a pine (Pinus caribaea Morelet) plantation in the Luquillo Experimental Forest in Puerto Rico. Soil surface CO₂ efflux was measured using the alkali trap method at 12 randomly-distributed locations in each treatment (control, root exclusion, litter exclusion, and both root and litter exclusion) in the plantation and the SF, respectively. We measured soil CO₂ efflux every two months and collected soil samples at each sampling location in different seasons to determine microbial biomass from August 1996 to July 1997. We found that soil CO₂ efflux was significantly reduced in the litter and root exclusion plots (7-year litter and/or root exclusion) in both the secondary forest and the pine plantation compared with the control. The reduction of soil CO₂ efflux was 35.6% greater in the root exclusion plots than in the litter exclusion plots in the plantation, whereas a reversed pattern was found in the secondary forest. Microbial biomass was also reduced during the litter and root exclusion period. In the root exclusion plots, total fungal biomass averaged 31.4% and 65.2% lower than the control plots in the plantation and the secondary forest, respectively, while the total bacterial biomass was 24% and 8.3% lower than the control plots in the plantation and the secondary forest, respectively. In the litter exclusion treatment, total fungal biomass averaged 69.2% and 69.7% lower than the control plots in the plantation and the secondary forest, respectively, while the total bacterial biomass was 48% and 50.1% lower than the control plots in the plantation and the secondary forest, respectively. Soil CO₂ efflux was positively correlated with both fungal and bacterial biomass in both the plantation the secondary forest. The correlation between soil CO₂ efflux and active fungal biomass was significantly higher in the plantation than in the secondary forest. However, the correlation between the soil CO₂ efflux and both the active and total bacterial biomass was significantly higher in the secondary forest than in the plantation in the day season. In addition, we found soil CO₂ efflux was highly related to the strong interactions among root, fungal and bacterial biomass by multiple regression analysis (R² > 0.61, P < 0.05). Our results suggest that carbon input from aboveground litterfall and roots (root litter and exudates) is critical to the soil microbial community and ecosystem carbon cycling in the wet tropical forests.     

Coupling of canopy gas exchange with root and rhizosphere respiration in a semi-arid forest

The strength of coupling between canopy gas exchange and root respiration was examined in ~15-yr-old ponderosa pine (Pinus ponderosa Doug. Ex Laws.) growing under seasonally drought stressed conditions. By regularly watering part of the root system to reduce tree water stress and measuring soil CO₂ efflux on the dry, distant side of the tree, we were able to determine the strength of the relationship between soil autotrophic (root and rhizosphere) respiration and changes in canopy carbon uptake and water loss by comparison with control trees (no watering). After ~40 days the soil CO₂ efflux rate, relative to pre-treatment conditions, was twice that of the controls. This difference, attributable to root and rhizosphere respiration, was strongly correlated with differences in transpiration rates between treatments (r² = 0.73, p<0.01). By the end of the period, transpiration of the irrigated treatment was twice that of controls. Periodic measurements of photosynthesis under non-light limited conditions paralleled the patterns of transpiration and were systematically higher in the irrigated treatment. We observed no evidence for a greater sensitivity of soil autotrophic respiration to temperature compared to the response of heterotrophic respiration to temperature; the Q₁₀ for total soil respiration was 1.6 (p>0.99) for both treatments. At the ecosystem scale, daily soil CO₂ efflux rate was linearly related to gross primary productivity (GPP) as measured by eddy-covariance technique (r² = 0.55, p<0.01), suggesting patterns of soil CO₂ release appear strongly correlated to recent carbon assimilation in this young pine stand. Collectively the observed relationships suggest some consideration should be given to the inclusion of canopy processes in future models of soil respiration.     

The effects of heating,rhizosphere, and depth on root litter decomposition are mediated by soil moisture

The breakdown and decomposition of plant inputs are critical for nutrient cycling, soil development, and climate-ecosystem feedbacks, but uncertainties persist in how the rates and products of litter decomposition are affected by soil temperature, rhizosphere, and depth of input. We investigated the effects of soil warming (+ 4 °C), rhizosphere, and depth of litter placement on the decomposition of Avena fatua (wild oat grass) root litter in a Mediterranean grassland ecosystem. Field lysimeters were subjected to three environmental treatments (heating, control, and plant removal) and three ¹³C-labeled root litter addition treatments (to A horizon, to B horizon, and no-addition disturbance control) for each of two harvest time points. We buried root litter in February 2014 and measured loss of ¹³C in CO₂ from the soil surface and in leachate as dissolved organic carbon (DOC) over two growing seasons. At the end of each growing season we recovered the ¹³C remaining in the soil. Loss of root litter C occurred almost entirely via heterotrophic respiration, with an estimated < 2% lost as DOC during the initial decay period. The added roots were broken down and incorporated into bulk soil material very quickly; only ~ 30% of added root was visible after 6 months. In the first growing season, decomposition occurred faster in the B than in the A horizon, the latter having greater moisture limitation. Subsequently, there was almost no further decomposition in the B horizon. After two growing seasons, less than 20% of the added root litter C remained in the A or B horizons of all environmental treatments. Heating did not stimulate decomposition, likely because it exacerbated the moisture limitation. However, while plots without plants dried down more slowly than plots with plants, their decomposition rate was not significantly greater, possibly due to the lack of rhizosphere processes such as priming. We conclude that in this Mediterranean grassland ecosystem, soil moisture, which is affected by season, depth, heating, and rhizosphere, plays a dominant role in mediating the effect of those factors on root litter decomposition, which after two seasons did not differ by depth or by treatment.