Soil respiration and carbon balance in a subtropical native forest and two managed plantations

From 1999 to 2003, a range of carbon fluxes was measured and integrated to establish a carbon balance for a natural evergreen forest of Castanopsis kawakamii (NF) and adjacent monoculture evergreen plantations of C. kawakamii (CK) and Chinese fir (Cunninghamia lanceolata, CF) in Sanming Nature Reserve, Fujian, China. Biomass carbon increment of aboveground parts and coarse roots were measured by the allometric method. Above- and belowground litter C inputs were assessed by litter traps and sequential cores, respectively. Soil respiration (SR) was determined by the alkaline absorbance method, and the contribution from roots, above- and belowground litters was separated by the DIRT plots. Annual SR averaged 13.742 t C ha⁻¹ a⁻¹ in the NF, 9.439 t C ha⁻¹ a⁻¹ in the CK, and 4.543 t C ha⁻¹ a⁻¹ in the CF. For all forests, SR generally peaked in later spring or early summer (May or June). The contribution of root respiration ranged from 47.8% in the NF to 40.3% in the CF. On average, soil heterotrophic respiration (HR) was evenly distributed between below- (47.3∼54.5%) and aboveground litter (45.5%–52.7%). Annual C inputs (t C ha⁻¹ a⁻¹) from litterfall and root turnover averaged 4.452 and 4.295, 4.548 and 2.313, and 2.220 and 1.265, respectively, in the NF, CK, and CF. As compared to HR, annual net primary production (NPP) of 11.228, 13.264, and 6.491 t C ha⁻¹ a⁻¹ in the NF, CK, and CF brought a positive net ecosystem production (NEP) of 4.144, 7.514, and 3.677 t C ha⁻¹ a⁻¹, respectively. It suggests that native forest in subtropical China currently acts as an important carbon sink just as the timber plantation does, and converting native forest to tree plantations locally during last decades might have caused a high landscape carbon loss to the atmosphere.     

Low stocks of coarse woody debris in a southwest Amazonian forest

The stocks and dynamics of coarse woody debris (CWD) are significant components of the carbon cycle within tropical forests. However, to date, there have been no reports of CWD stocks and fluxes from the approximately 1.3 million km² of lowland western Amazonian forests. Here, we present estimates of CWD stocks and annual CWD inputs from forests in southern Peru. Total stocks were low compared to other tropical forest sites, whether estimated by line-intercept sampling (24.4 ± 5.3 Mg ha⁻¹) or by complete inventories within 11 permanent plots (17.7 ± 2.4 Mg ha⁻¹). However, annual inputs, estimated from long-term data on tree mortality rates in the same plots, were similar to other studies (3.8 ± 0.2 or 2.9 ± 0.2 Mg ha⁻¹ year⁻¹, depending on the equation used to estimate biomass). Assuming the CWD pool is at steady state, the turnover time of coarse woody debris is low (4.7 ± 2.6 or 6.1 ± 2.6 years). These results indicate that these sites have not experienced a recent, large-scale disturbance event and emphasise the distinctive, rapid nature of carbon cycling in these western Amazonian forests.     

Effects of elevated CO2 on growth and carbon/nutrient balance in the deciduous woody shrub Lindera benzoin (L.) Blume (Lauraceae)

We examined the effects of elevated CO₂ on growth and carbon/nutrient balance in a natural population of the deciduous temperate zone shrub Lindera benzoin. Our data concern whole plant, leaf, and stem growth for the first two seasons of a long-term field experiment in which CO₂ levels were manipulated in situ. In addition to growth parameters, we evaluated changes in leaf and stem chemistry, including total nitrogen, nonstructural carbohydrates, and total phenolics. Over the course of this study, L. benzoin appeared to respond to elevated CO₂ primarily by physiological and biochemical changes, with only a slight enhancement in aboveground growth (ramet height). Positive effects on aboveground growth were primarily evident in young (nonreproductive) ramets. Our results suggest that nitrogen limitation may have constrained plants to allocate carbohydrates produced in response to elevated CO₂ primarily to storage and belowground growth, and perhaps to increased secondary chemical production, rather than to increased stem and leaf growth. We discuss our results in terms of changes in carbon/nutrient balance induced by elevated CO₂, and provide predictions for future changes in this system based upon constraints imposed by intrinsic and extrinsic factors and their potential effects on the reallocation of stored reserves.     

Quantitative analysis of carbon balance in the reproductive organs and leaves of <Emphasis Type="Italic">Cinnamomum camphora</Emphasis> (L.) Presl

We investigated seasonal changes in dry mass and CO₂ exchange rate in fruit and leaves of the evergreen tree Cinnamomum camphora with the aim of quantitatively determining the translocation balance between the two organs. The fruit dry mass growth peaked in both August and October: the first increase was due to fruit pulp development and the second to seed development. Fruit respiration also increased with the rapid increase in fruit dry mass. Therefore, the carbohydrates required for fruit development showed two peaks during the reproductive period. Fruit photosynthesis was relatively high in early August, when fruit potentially re-fixed 75% of respired CO₂, indicating that fruit photosynthesis contributed 15–35% of the carbon requirement for fruit respiration. Current-year leaves completed their growth in June when fruit growth began. Current-year leaves translocated carbohydrates at a rate of approximately 10–25 mg dry weight (dw) leaf⁻¹ day⁻¹ into other organs throughout the entire fruit growth period. This rate of translocation from current-year leaves was much higher than the amount of carbohydrate required for reproduction (ca. 3 mg dw fruit⁻¹ day⁻¹). Given the carbon balance between fruit and current-year leaves, carbohydrates for reproduction were produced within the current-year fruit-bearing shoots. C. camphora would be adaptive for steadily supplying enough amount of carbohydrate to the fruits, as there was little competition for carbohydrates between the two organs. As assimilates by leaves are used for processes such as reproduction and the formation of new shoots, photosynthesis by reproductive organs is considered to be important to compensate for reproductive cost.     

Simulated annual carbon fluxes of grassland ecosystems in extremely arid conditions

In order to understand how changes in climate and land cover affect carbon cycles and structure and function of regional grassland ecosystems, we developed a grassland landscape productivity model, proposed an approach that combined both process-based modeling and spatial analysis with field measurements, and provided an example of semiarid region in Inner Mongolia, China, in extremely arid conditions. The modeled monthly mean and total net primary productivity, and monthly and annual mean respiration rates for Leymus chinensis steppes during the growing seasons in 2002 were mostly within the normal varying ranges of measured values, or similar to the field measurements, conducted in the similarly arid conditions. And the modeled total net ecosystem productivity (NEP) for L. chinensis steppes and Stipa grandis steppes were both between the lower and the higher measurements or within modeled multi-annual data by the other model. The modeled total NEP was 1.91 g C/m²/year over the entire study region. It indicated that if human disturbances were not considered, carbon budget over the entire study region during the growing seasons was nearly in balance or weak carbon sink even under extremely arid conditions. However, the modeled NEP spatially greatly varied not only over the entire study region (−48.28–52.09 g C/m²/year), but also among different land cover types. The modeled results also showed that there were obvious seasonal variations in carbon fluxes, mainly caused by leaf area index; and annual precipitation was the major limiting factor for the obvious spatial patterns of carbon fluxes from the east to the west. The modeled results also revealed the influence of extreme drought on carbon fluxes. The study provides an effective approach to derive useful information about carbon fluxes for different land cover types, and thus can instruct regional land-use planning and resource management based on carbon storage conditions.     

A carbon balance model of the sorghum-Striga hermonthica host-parasite association

Abstract. Growth and gas exchange measurements are used to formulate a carbon balance model to describe the sorghum- Striga hermonthica host-Parasite association. S. hermonthica reduces the growth and radically alters the architecture of infected sorghum plants. Grain and stem weight are reduced, whilst leaf and root biomass are maintained. Losses in host productivity result from two processes: export of carbon to the parasite and Parasite-induced reductions in host photosynthesis. The latter occurs before the emergence of the Parasite above ground and accounts for 80% of the Predicted loss in production over the lifecycle of the association. S. hermonthica is dependent on carbon exported from the host, since the plant has low rates of photosynthesis coupled with high rates of respiration. Host-derived carbon accounts for approximately one-third of the total parasite carbon requirement.     

Pollination and breeding systems of woody plant species in tropical dry evergreen forests,southern India

Tropical dry evergreen forests (TDEF) are a unique forest type found along the east coast of India. They mostly occur as small, isolated fragments of varying sizes (0.5 to ≈10 ha) and are considered as endangered forests types in peninsular India. Although plant diversity is well documented in these forests, there is a paucity of ecological studies vital for conservation and for planning restoration activities. We studied reproductive biology of 13 woody species: four trees, six shrubs, and three lianas in fragments of TDEF in southern India. The phenology of reproduction, floral biology, anthesis and sexual system of each species were recorded. The pollination mode was assessed through observations of the visitation frequency of pollinators and from the floral characters. The breeding system was determined by hand-augmented self- and cross-pollination experiments. The plants flowered during the dry season from January to July. Plants of nine species had both flowers and fruits at the same time. Twelve species were hermaphrodites and one was polygamo-dioecious. Flowers of 11 species opened at dawn and two at dusk. Four species were self-incompatible and six were self-compatible. Natural fruit set ranged from 10% to 56%, self-incompatible species having low fruit set. Cross-pollen augmentation increased fruit set, suggesting presence of outcrossing in all species. The majority of plants species (85%) had a generalized pollination system, receiving visits from diverse insects, such as social bees, solitary bees, wasps, moths and flies. However, only few of them were functionally important for the species. Two species namely: Capparis brevispina and C. zeylanica had butterflies and birds, respectively, as their main pollinators. Our data reveal that there is a predominance of outcrossing in plant species and a generalized pollination system in these forests. We suggest that restoration of TDEFs is crucial as habitats, not only for wild plants but also for pollinating insects.     

Translocation and utilization of carbon in wheat (Triticum aestivum)

Wheat ( Triticum aestivum L. cv. SUN 9E) was grown in a growth chamber under conditions of low soil nitrogen. Translocation of carbon to the roots and the subsequent utilization of these carbohydrates was determined. In vegetative plants (22 days old), 21.5 mg C day⁻¹ were translocated to the roots. 29% of this was incorporated into dry matter, 32% was respired (28% via the cytochrome and 4% via a SHAM-sensitive, presumably the alternative nonphosphorylating, pathway) and 39% was translocated back to the shoots, mainly in the form of amino acids. – The rote of root maintenance respiration during the vegetative phase was estimated to be 0.7 mg O₂ h⁻¹ (g dry weight of roots)⁻¹ and the root growth respiration to be 0.41 g O₂ (g dry weight of roots)⁻¹. Total carbohydrate utilization due to root respiration via the alternative, nonphosphorylating pathway during the major part of the growth period was calculated to be only ca 6% of carbohydrate utilization for grain growth. The rate of specific mass transfer (SMT) of sugars in the sieve tubes was estimated from the data on C-translocation and data on the total area occupied by sieve tubes in a cross section of the root system. SMT was calculated to be 0.8 mg sucrose s⁻¹ cm⁻², which is very similar to the published value on SMT for other organs, except roots.     

Responses of dry weight growth under SO2 stress in an SO2-tolerant plant,Polygonum cuspidatum

The effects of exposure to 0.5–0.7 ppm SO₂ for about one month on the dry weight growth and net photosynthesis ofPolygonum cuspidatum were investigate. Furthermore, the carbon and nitrogen concentrations in each plant organ were measured. The results obtained showed no significant decrease in the total dry weight of SO₂-treated plants in comparison with controls. On the other hand, the leaf area (LA) and/or leaf dry weight of SO₂-treated plants were increased, and the root dry weight (RW) was decreased, in comparison with controls. The leaf carbon assimilation rate (CAR) in SO₂-treated plants was slightly decreased in spite of a clear decrease in net photosynthesis, and the value of (SW+RW)/LA (SW stem dry weight) was decreased in comparison with controls, thus minimizing the reduction in CAR. Furthermore, the ratio of total leaf carbon absorption (leaf area x CAR) to total root nitrogen absorption (root dry weight x nitrogen assimilation rate) in SO₂-treated plants was similar to that in controls. From these results, it can be concluded that an increase in leaf area and/or leaf dry weight and a decrease in root dry weight inP. cuspidatum under SO₂ stress may be induced in order to compensate for the decrease in CAR and to maintain the ratio of total leaf carbon absorption to total root nitrogen absorption in the early stage of vegetative growth.     

Retranslocation of carbon reserves from the woody storage tissues into the fruit as a response to defoliation stress during the ripening period in Vitis vinifera L.

A technique for reliable labeling of the carbon reserves of the trunk and roots without labeling the current year's growth of grapevines was developed in order to study retranslocation of carbon from the perennial storage tissues into the fruit in response to defoliation stress during the ripening period. A special training system with two shoots was used: the lower one (feeding shoot) was cut back and defoliated to one single leaf (¹⁴CO₂-feeding leaf) while the other (main shoot) was topped to 12 leaves. The potted plants were placed in a water bath at 30 °C to increase root temperature and therefore their sink activity. Additionally, a cold barrier (2–4 °C) was installed at the base of the main shoot to inhibit acropetal ¹⁴C translocation. Using this method, we were able to direct labeled assimilates to trunk and roots in preference to the current year's growth. On vines with root and shoot at ambient temperature, 44% of the ¹⁴C activity was found in the main shoot 16 h after feeding whereas only 2% was found in the temperature-treated vines. At the onset of fruit ripening, and at three-week intervals thereafter until harvest, potted grapevines were fed with ¹⁴CO₂ using the temperature treatment described above. Sixteen hours after feeding, half of the vines of each group were defoliated by removing all except the two uppermost main leaves. Three weeks after each treatment, vines were destructively harvested and the dry weight and ¹⁴C incorporation determined for all plant parts. Under non-stressing conditions, there was no retranslocation of carbon reserves to support fruit maturation. Vines responded to defoliation stress by altering the natural translocation pattern and directing carbon stored in the lower parts to the fruit. In the three weeks following veraison (the inception of ripening in the grape berry), 12% of the labeled carbon reserves was translocated to the fruit of defoliated plants compared to 1.6% found in the clusters of control vines. Retranslocation from trunk and roots was highest during the middle of the ripening period, when 32% of the labeled carbon was found in the fruit compared to 0.7% in control plants. Defoliation during this period also caused major changes in dry-matter partitioning: the fruit represented 31% of total plant biomass compared to 21% measured in the control vines. Root growth was reduced by defoliation at veraison and during the ripening period. Defoliation three weeks before harvest did not affect dry matter or ¹⁴C partitioning.     

Seasonal patterns of canopy development and carbon gain in nineteen warm desert shrub species

Summary Canopy development and photosynthetic rate were measured at monthly intervals over a period of one year in 19 shrub and subshrub species of the Mojave and upper Sonoran Deserts. Thirteen of these species realized a substantial fraction of their total net carbon assimilation via twig photosynthesis. The twig contribution to whole plant yearly carbon gain reached a maximum of 83% in species such as Thamnosma montana, Salizaria mexicana, and Baccharis brachyphylla. This large contribution by twigs was due to both low levels of leaf production and the greater longevity of twig tissues. In some other species, however, leaf and twig organs had similar lifespans. During the year of this study (which had an unusually warm, mild winter), no species showed a pattern of winter deciduousness. The reduction in total photosynthetic area between maximal spring canopy development and mid August summer dormancy ranged from 32 to 94%. Some herbaceous perennial species died back to the ground, but none of the woody shrubs were totally without green canopy area at any time of the year. No species studied were capable of high rates of photosynthesis at low plant water potentials in July and August, but, in those species which maintained a substantial canopy area through the drought period, previously stressed tissues showed substantial recovery after fall rains. Photosynthetic rate was significantly correlated with both plant water potential and tissue nitrogen content over the entire year, but only weakly so. This is due in part to the winter months when plant water potentials and tissue nitrogen contents were high, but photosynthetic rates were often low.     

Assimilation,respiration and allocation of carbon inPlantago major as affected by atmospheric CO2 levels

The response ofPlantago major ssp,pleiosperma plants, grown on nutrient solution in a climate chamber, to a doubling of the ambient atmospheric CO₂ concentration was investigated. Total dry matter production was increased by 30% after 3 weeks of exposure, due to a transient stimulation of the relative growth rate (RGR) during the first 10 days. Thereafter RGR returned to the level of control plants. Photosynthesis, expressed per unit leaf area, was stimulated during the first two weeks of the experiment, thereafter it dropped and nearly reached the level of the control plants. Root respiration was not affected by increased atmospheric CO₂ levels, whereas shoot, dark respiration was stimulated throughout the experimental period. Dry matter allocation over leaves stems and roots was not affected by the CO₂ level. SLA was reduced by 10%, which can partly be explained by an increased dry matter content of the leaves. Both in the early and later stages of the experiment, shoot respiration accounted for a larger part of the carbon budget in plants grown at elevated atmospheric CO₂. Shifts in the total carbon budget were mainly due to the effects on shoot respiration. Leaf growth accounted for nearly 50% of the C budget at all stages of the experiment and in both treatments.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - RGR relative growth rate - R/S root to shoot ratio - RWR root weight ratio - SLA specific leaf area - SWR stem weight ratio     

Respiratory carbon use and carbon storage in mid‐rotation loblolly pine (Pinus taeda L.) plantations: the effect of site resources on the stand carbon balance

We used estimates of autotrophic respiration (R_A), net primary productivity (NPP) and soil CO₂ evolution (S_ff), to develop component carbon budgets for 12‐year‐old loblolly pine plantations during the fifth year of a fertilization and irrigation experiment. Annual carbon use in R_A was 7.5, 9.0, 15.0, and 15.1 Mg C ha^?1 in control (C), irrigated (I), fertilized (F) and irrigated and fertilized (IF) treatments, respectively. Foliage, fine root and perennial woody tissue (stem, branch, coarse and taproot) respiration accounted for, respectively, 37%, 24%, and 39% of R_A in C and I treatments and 38%, 12% and 50% of R_A in F and IF treatments. Annual gross primary production (GPP=NPP+R_A) ranged from 13.1 to 26.6 Mg C ha^?1. The I, F, and IF treatments resulted in a 21, 94, and 103% increase in GPP, respectively, compared to the C treatment. Despite large treatment differences in NPP, R_A, and carbon allocation, carbon use efficiency (CUE=NPP/GPP) averaged 0.42 and was unaffected by manipulating site resources. Ecosystem respiration (R_E), the sum of S_ff, and above ground R_A, ranged from 12.8 to 20.2 Mg C ha^?1 yr^?1. S_ff contributed the largest proportion of R_E, but the relative importance of S_ff decreased from 0.63 in C treatments to 0.47 in IF treatments because of increased aboveground R_A. Aboveground woody tissue R_A was 15% of R_E in C and I treatments compared to 25% of R_E in F and IF treatments. Net ecosystem productivity (NEP=GPP‐R_E) was roughly 0 in the C and I treatments and 6.4 Mg C ha^?1 yr^?1 in F and IF treatments, indicating that non‐fertilized treatments were neither a source nor a sink for atmospheric carbon while fertilized treatments were carbon sinks. In these young stands, NEP is tightly linked to NPP; increased ecosystem carbon storage results mainly from an increase in foliage and perennial woody biomass.     

Root respiration and its importance for the carbon balance of beech saplings (Fagus sylvatica L.) in a montane beech forest

Root respiration of 10-year-old beech saplings (Fagus sylvatica L.) grown in the understorey (UND) and in a natural gap (GAP) of a mature beech forest in the Solling mountains, FRG, was investigated from April until December, 1990. Respiration rates of fine, medium and coarse roots were measured in situ by a PC-controlled cuvette system. Fine root respiration rates were in the range of 0.5–9.8 nmol CO₂ gDW^–1 s^–1 at both sites, but respiration rates of UND saplings were higher, compared to those of GAP saplings. The dependence of respiratory activity on soil temperature proved to be highly significant (p<0.001) for both plots, following a quasi-Arrhenius type curve. Fine root respiration rates of UND saplings were highly significantly, negatively correlated with the water content of the attached organic material, whereas respiration rates of GAP saplings did not show such a correlation. Further, a significant correlation (p<0.01) between mycorrhizal biomass and respiration rate was detected at the UND site, but not at the GAP site. Medium and coarse root respiration rates were very similar and no significant differences between the two sites were detected. Maximum respiration rates of 3.1 nmol CO₂ gDW^–1 s^–1 were reached in the middle of July. Due to low light intensities in the under storey, daily net CO₂ assimilation rates of UND saplings were much smaller than those of GAP saplings. At both sites, net CO₂ assimilation rates varied more than respiration rates and thus the carbon balance of beech saplings was more affected by the rate of carbon fixation than by the rate of respiratory carbon loss.     

Resource allocation to reproduction and soma in Drosophila: a stable isotope analysis of carbon from dietary sugar

Metabolic resources in adults of holometabolous insects may derive either from larval or adult feeding. In Drosophila melanogaster, reproduction and lifespan are differently affected by larval vs. adult resource availability, and it is unknown how larval vs. adult acquired nutrients are differentially allocated to somatic and reproductive function. Here we describe the allocation of carbon derived from dietary sugar in aging female D. melanogaster. Larval and adult flies were fed diets contrasting in sucrose (13)C/(12)C, from which we determined the extent to which carbon acquired at each stage contributed to adult somatic tissue and to egg manufacture. Dietary sugar is very important in egg provisioning; at every age, roughly one half of the carbon in eggs was derived from sugar, which turned over from predominantly larval to entirely adult dietary sources. Sucrose provided approximately 40% of total somatic carbon, of which adult dietary sucrose came to supply approximately 75%. Unlike in eggs, however, adult acquired sucrose did not entirely replace the somatic carbon from larvally acquired sucrose. Because carbon from larval sucrose appears to be fairly "replaceable", larval sucrose cannot be a limiting substrate in resource allocation between reproduction and lifespan.     

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.