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Incidence of agonal respirations in sudden cardiac arrest.

STUDY OBJECTIVE: To discover the frequency of agonal respirations in cardiac arrest calls, the ways callers describe them, and discharge rates associated with agonal respirations. DESIGN: We reviewed taped recordings of calls reporting cardiac arrests and emergency medical technician and paramedic incident reports for 1991. Arrests after arrival of emergency medical services were excluded. SETTING: King County, Washington, excluding the city of Seattle. PARTICIPANTS: Four hundred forty-five persons with out-of-hospital cardiac arrests receiving emergency medical services. INTERVENTIONS: Telephone CPR, emergency medical technicians-defibrillation, and advanced life support by paramedics. MEASUREMENTS AND MAIN RESULTS: Any attempts at breathing described by callers were identified, as well as whether agonal respirations could be heard by dispatcher, emergency medical technicians, or paramedics. Agonal respirations occurred in 40% of 445 out-of-hospital cardiac arrests. Callers described agonal breathing in a variety of ways. Agonal respirations were present in 46% of arrests caused by cardiac etiology compared with 32% in other etiologies (P < .01). Fifty-five percent of witnessed arrests had agonal activity compared with 16% of unwitnessed arrests (P < .001). Agonal respirations occurred in 56% of arrests with a rhythm of ventricular fibrillation compared with 34% of cases with a nonventricular fibrillation rhythm (P < .001). Twenty-seven percent of patients with agonal respirations were discharged alive compared with 9% without them (P < .001). CONCLUSION: There is a high incidence of agonal activity associated with out-of-hospital cardiac arrest. Presence of agonal respirations is associated with increased survival. These findings have implications for public CPR training programs and emergency dispatcher telephone CPR programs.

Emergency Medical Services↗

4-Hydroxy-2(E)-nonenal inhibits CNS mitochondrial respiration at multiple sites.

A destructive cycle of oxidative stress and mitochondrial dysfunction is proposed in neurodegenerative disease. Lipid peroxidation, one outcome of oxidative challenge, can lead to the formation of 4-hydroxy-2(E)-nonenal (HNE), a lipophilic alkenal that forms stable adducts on mitochondrial proteins. In this study, we characterized the effects of HNE on brain mitochondrial respiration. We used whole rat brain mitochondria and concentrations of HNE comparable to those measured in patients with Alzheimer's disease. Our results showed that HNE inhibited respiration at multiple sites. Complex I-linked and complex II-linked state 3 respirations were inhibited by HNE with IC50 values of approximately 200 microM HNE. Respiration was apparently diminished owing to the inhibition of complex III activity. In addition, complex II activity was reduced slightly. The lipophilicity and adduction characteristics of HNE were responsible for the effects of HNE on respiration. The inhibition of respiration was not prevented by N-acetylcysteine or aminoguanidine. Studies using mitochondria isolated from porcine cerebral cortex also demonstrated an inhibition of complex I- and complex II-linked respiration. Thus, in neurodegenerative disease, oxidative stress may impair mitochondrial respiration through the production of HNE.

Aldehydes↗

Impact of needle age on the response of respiration in Scots pine to long-term elevation of carbon dioxide concentration and temperature.

Sixteen 20-year-old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed in environment-controlled chambers that for 4 years maintained: (1) ambient conditions (CON); (2) elevated atmospheric carbon dioxide concentration [CO2] (ambient + 350 micromol mol-1; EC); (3) elevated temperature (ambient + 2-3 degrees C; ET); or (4) elevated [CO2] and temperature (EC+ET). Dark respiration rate, specific leaf area (SLA) and the concentrations of starch and soluble sugars in needles were measured in the fourth year. Respiration rates, on both an area and a mass basis, and SLA decreased in EC relative to CON, but increased in ET and EC+ET, regardless of needle age class. Starch and soluble sugar concentrations for a given needle age class increased in EC, but decreased slightly in ET and EC+ET. Respiration rates and SLA were highest in current-year needles in all treatments, whereas starch and soluble sugar concentrations were highest in 1-year-old needles. Relative to that of older needles, respiration of current-year needles was inhibited less by EC, but increased in response to ET and EC+ET. All treatments enhanced the difference in respiration between current-year and older needles relative to that in CON. Age had a greater effect on needle respiration than any of the treatments. There were no differences in carbohydrate concentration or SLA between needle age classes in response to any treatment. Relative to CON, the temperature coefficient (Q10) of respiration increased slightly in EC, regardless of age, but declined significantly in ET and EC+ET, indicating acclimation of respiration to temperature.

Carbohydrate Metabolism↗

Spatial and temporal variations in soil respiration in relation to stand structure and soil parameters in an unmanaged beech forest.

Soil CO2 efflux (soil respiration) plays a crucial role in the global carbon cycle and efflux rates may be strongly altered by climate change. We investigated the spatial patterns of soil respiration rates in 144 measurement locations in a 0.5-ha plot and the temporal patterns along a 300-m transect in the 0.5-ha plot. Measurements were made in an unmanaged, highly heterogeneous beech forest during 2000 and 2001. We investigated the effects of soil, roots and forest stand structure on soil respiration, and we also assessed the stability of these spatial patterns over time. Soil temperature alone explained between 68 and 95% of the temporal variation in soil respiration; however, pronounced spatial scatter of respiration rates was not explained by soil temperature. The observed spatial patterns stayed remarkably stable throughout the growing season and over 2 years. The most important structural parameter of the stand was the mean diameter at breast height of trees within a distance of 4 m of the measurement locations (m-dbh4), which explained 10-19% of the variation in soil respiration throughout the growing season. Among the soil chemical parameters, carbon content (bulk as well as dissolved) and magnesium content explained 62% of the spatial variation in soil respiration. The final best model combining soil, root and stand structural parameters (fine root biomass, soil carbon content, m-dbh4 and soil water content) explained 79% of the variation in soil respiration, illustrating the importance of both biotic and abiotic factors.

Carbon Dioxide↗

Fine root respiration in the mangrove Rhizophora mangle over variation in forest stature and nutrient availability.

Root respiration uses a significant proportion of photosynthetically fixed carbon (C) and is a globally important source of C liberated from soils. Mangroves, which are an important and productive forest resource in many tropical and subtropical countries, sustain a high ratio of root to shoot biomass which may indicate that root respiration is a particularly important component in mangrove forest carbon budgets. Mangroves are often exposed to nutrient pollution from coastal waters. Here we assessed the magnitude of fine root respiration in mangrove forests in Belize and investigated how root respiration is influenced by nutrient additions. Respiration rates of excised fine roots of the mangrove, Rhizophora mangle L., were low (4.01 +/- 0.16 nmol CO(2) g(-1) s(-1)) compared to those measured in temperate tree species at similar temperatures. In an experiment where trees where fertilized with nitrogen (N) or phosphorus (P) in low productivity dwarf forests (1-2 m height) and more productive, taller (4- 7 m height) seaward fringing forests, respiration of fine roots did not vary consistently with fertilization treatments or with forest stature. Fine roots of taller fringe trees had higher concentrations of both N and P compared to dwarf trees. Fertilization with P enhanced fine root P concentrations in both dwarf and fringe trees, but reduced root N concentrations compared to controls. Fertilization with N had no effect on root N or P concentrations. Unlike photosynthetic C gain and growth, which is strongly limited by P availability in dwarf forests at this site, fine root respiration (expressed on a mass basis) was variable, but showed no significant enhancements with nutrient additions. Variation in fine root production and standing biomass are, therefore, likely to be more important factors determining C efflux from mangrove sediments than variations in fine root respiration per unit mass.

Cell Respiration↗

Respiration metabolism reduces oxidative and acid stress to improve long-term survival of Lactococcus lactis.

The impact of oxygen on a cell is strongly dependent on its metabolic state: survival in oxygen of free-living Lactococcus lactis, best known as a fermenting, acidifying bacterium, is generally poor. In contrast, if haem is present, L. lactis uses oxygen to switch from fermentation to respiration metabolism late in growth, resulting in spectacularly improved long-term survival. Oxygen is thus beneficial rather than detrimental for survival if haem is provided. We examined the effects of respiration on oxygen toxicity by comparing integrity of stationary phase cells after aerated growth without and with added haem. Aeration (no haem) growth caused considerable cellular protein and chromosomal DNA damage, increased spontaneous mutation frequencies and poor survival of recA mutants. These phenotypes were greatly diminished when haem was present, indicating that respiration constitutes an efficient barrier against oxidative stress. Using the green fluorescent protein as an indicator of intracellular oxidation state, we showed that aeration growth provokes significantly greater oxidation than respiration growth. Iron was identified as a main contributor to mortality and DNA degradation in aeration growth. Our results point to two features of respiration growth in lactococci that are responsible for maintaining low oxidative damage: One is a more reduced intracellular state, which is because of efficient oxygen elimination by respiration. The other is a higher pH resulting from the shift from acid-forming fermentation to respiration metabolism. These results have relevance to other bacteria whose respiration capacity depends on addition of exogenous haem.

Bacterial Proteins↗

Influence of Cheyne-Stokes respiration on cardiovascular oscillations in heart failure.

In patients with congestive heart failure, Cheyne-Stokes respiration is accompanied by oscillations in blood pressure and heart rate at a very low frequency. It is not known whether these cardiovascular oscillations are primarily related to oscillations in ventilation or oxyhemoglobin saturation. We hypothesized that abolition of the ventilatory oscillations of Cheyne-Stokes respiration by CO2 inhalation would eliminate accompanying oscillations in blood pressure and heart rate but that elimination of hypoxic dips by supplemental O2 would not. We studied 10 subjects with heart failure and Cheyne-Stokes respiration during sleep using frequency spectral analysis. During Cheyne-Stokes respiration, heart rate and blood pressure oscillated in association with respiratory oscillations at very low frequency. Inhalation of CO2 abolished Cheyne-Stokes respiration and associated oscillations in both blood pressure and heart rate. In contrast, inhalation of O2 sufficient to eliminate hypoxic dips had no significant effect on Cheyne-Stokes respiration, blood pressure (n = 6), or heart rate (n = 5). We conclude that ventilatory oscillations during Cheyne-Stokes respiration rather than oscillations in oxygenation per se powerfully induce heart rate and blood pressure oscillations. Cheyne-Stokes respiration is therefore one of the mechanisms that contributes to the very low-frequency oscillations in heart rate and blood pressure observed in patients with heart failure.

Administration, Inhalation↗

Changes in heart rate during feedback control of respiration.

Subjects received one of eight treatments: practice at increasing heart rate with heart-rate feedback, practice at decreasing heart rate with heart-rate feedback, practice at increasing heart rate without heart-rate feedback, practice at decreasing heart rate without heart-rate feedback, practice at increasing respiration rate with respiratory feedback, practice at decreasing respiration rate with respiratory feedback, practice at increasing respiration rate with respiratory instructions only, practice at decreasing respiration rate with respiratory instructions only. Heart rate, Respiration rate, and Respiration depth were measured. Analysis indicated that subjects who controlled respiration with respiratory feedback reliably increased and decreased heart rate; subjects who controlled respiration with respiratory instructions only reliably increased but not decreased heart rate; subjects in the respiratory-feedback conditions showed higher heart-rate increase and decrease than heart-rate increase and decrease of subjects in the other six conditions.

Adult↗

[A comparative study on soil respiration between grazing and fenced typical Leymus chinensis steppe, Inner Mongolia].

With enclosed chamber Method, this paper studied the soil respiration in grazing and fenced typical Leymus chinensis steppes, Inner Mongolia, and its relationships with environmental factors. The results showed that the daily pattern of soil respiration could be expressed as a one-humped curve, and the highest values appeared at 13:00-15:00 in the fenced and grazing plots. The diurnal dynamics of soil respiration mainly depended on the surface temperature at the fenced plots and the soil temperature at 5 cm depth at the grazing plots. In June and July, the average soil respiration rate was 2.7 times greater at the fenced plots than that at the grazing plots, while the difference was not distinct in August and September, which was similar with the change of the belowground biomass. The reason was probably that the plant was influenced differently in different phenological phases by grazing and the change of environmental factors. It showed that human activity may not result in the increase of soil respiration rate. The seasonal dynamics of soil respiration was closely correlated with soil water content at the 0-10 cm depth at the fenced and grazing sites, and the maximum R2 was 0.853 and 0.741, respectively. The difference was that the correlation of soil respiration seasonal dynamics with soil water content was larger at the fenced plots than at the grazing plots. The correlations of soil respiration diurnal and seasonal dynamics with temperature and soil water content at lower profiles were larger than those at deeper profiles at the fenced and grazing sites.

Animal Husbandry↗

[Soil respiration and its influencing factors at grazing and fenced typical Leymus chinensis Steppe, Nei Monggol].

Based on two-year soil respiration observing data at fenced and grazing typical Leymus chinensis steppe, Nei Monggol by the enclosed chamber method, soil respiration and its relationships with various affecting factors were analyzed. The major results include: (1) The diurnal and seasonal dynamics of soil respiration are similar at fenced and grazing typical Leymus chinensis steppe, but soil respiration rate is larger at the fenced plots than that at the grazing plots, and their values were 219.18 mg x (m2 x h)(-1) and 111.27 mg x (m2 x h)(-1), respectively. The reason of soil respiration rate increasing in the fenced plots might be the increases of biomass in the belowground and soil water content. (2) Among the influencing factors on soil respiration, grazing decreases the effects of soil water content and relative humidity and increases the effect of photosynthetic rate, and has no significant effects on other factors. (3) At the fenced plots, the correlation of soil respiration rate with soil water content is largest, followed by photosynthetic rate, air temperature, relative humidity, air CO2 concentration, intercellular CO2 concentration, stomatal conductance and the lowest being photosynthetically active radiation. Soil water content and air temperature are the main environmental factors, and photosynthetic rate is the critical biotic factor. The affecting factors on soil respiration are the same at the fenced and grazing plots, however the importance of those influencing factors is changed quantitatively.

Animals↗

[Relation between long-term artificial respiration and respiratory tract infection in 104 patients in a surgical intensive care unit].

The authors studied over a period of two years prospective connections between long-term artificial respiration and infections of the lower respiratory tract. A distinction was made between colonisation (RTC) and infection (RTI). Besides the incidence rate of RTC and RTI the severity of the infection was recorded. The connections between the underlying disease, the duration of artificial respiration, the age of the patient and the antibiotic treatment with the pathogenesis of RTC and RTI were studied. In more than half of the patients RTC could be demonstrated already within the first 24 hours; differences were related to underlying disease, start of ventilation and antibiotic treatment. The RTI incidence rate in all 104 ventilated patients was 67.3%. 19.2% of these already had RTI when respiration was initiated. Thus 48.1% of the patients developed RTI during artificial respiration. The diagnosis RTI was most frequent on the fourth day of respiration, and 80% of all RTI cases had appeared by the fifth day. The highest incidence was seen in patients with a thoracic trauma (85.7%). This group of patients also included the majority of life threatening cases of RTI. In one-quarter of the patients who developed RTI during artificial respiration there was no significant deterioration of pulmonary gas exchange; in 56% however, respiration was impaired to a life-threatening extent. The age of the patients did not have any bearing on the incidence rate and severity of RTI; however, all patients under artificial respiration who were over 70 years of age, died if the severity of the RTI had to be classified as life-threatening.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Nocturnal dyspnea and atrial fibrillation predict Cheyne-Stokes respirations in patients with congestive heart failure.

BACKGROUND: Cheyne-Stokes respirations have frequently been noted in highly selected groups of patients with congestive heart failure, but their prevalence in an unselected population with congestive heart failure is undefined. METHODS: One hundred consecutive unselected outpatients or stable inpatients with clinical congestive heart failure encountered by three clinical cardiologists during a 6-month period were screened for Cheyne-Stokes respirations with overnight oximetry. RESULTS: The mean age (+/- SD) of the patients was 70 +/- 8.6 years. Of the 100 patients, 33% had had previous coronary bypass surgery, 77% were men, 57% had hypertension, and 32% had atrial fibrillation. The mean ejection fraction (+/- SD) was 34% +/- 13%. Periodic breathing was assessed qualitatively as Cheyne-Stokes respirations in 27% of patients, nonspecific sleep-disordered breathing (apneas and/or hypopneas) in 43%, and normal in 30%. For patients with Cheyne-Stokes respirations, patients with nonspecific sleep-disordered breathing, and normal subjects, the mean numbers of oxyhemoglobin desaturation events per hour were 24, 10, and 2, and the total numbers of desaturations of 4% or more that lasted less than 3 minutes were 172, 74, and 13. Independent predictors of Cheyne-Stokes respirations vs non-Cheyne-Stokes respirations included a history of nocturnal dyspnea (odds ratio, 4.00; 95% confidence interval, 1.33 to 12.04; P = .01) and atrial fibrillation (odds ratio, 3.24; 95% confidence interval, 1.21 to 8.48; P = .02). CONCLUSIONS: Cheyne-Stokes respirations and nonspecific sleep-disordered breathing are common in unselected patients with congestive heart failure, and Cheyne-Stokes respirations are predicted by a history of nocturnal dyspnea and the presence of atrial fibrillation. Techniques designed to modify the nocturnal breathing pattern of patients with congestive heart failure may be applicable to a large portion of the congestive heart failure population.

Aged↗

Medical evaluation for respirator use.

The purpose of a respirator is to prevent the inhalation of harmful airborne substances or to provide a source of respirable air when breathing in oxygen-deficient atmospheres. For a physician to recommend the use of respirator, general background information on respiratory-protective devices is required. The first part of this clinical practice review describes the general aspects of industrial hygiene, respirators and a respirator-certification program. The second part addresses matters related to medical certification for respirator use. Medical certification for respirators is an important part of the activities of the occupational physician. To determine whether a worker is able to tolerate the added strain of a respiratory protective device is a complex process in which factors such as fitness for work, health of the individual, characteristics of the work itself, and the properties, type, and requirements of the respiratory protective device, have to be considered. Medical certification is of utmost importance for respirator use, and it should be viewed as an element in a comprehensive respiratory protection program. A comprehensive program is the key element in affording the workers' effective respiratory protection once the initial steps of the hierarchy of methods of hazard control have proved insufficient or infeasible. As a result, the need for the industrial hygiene/safety officer, the worker, the employer and the medical professional to work as a team is much more than in any other field of occupational medicine--a necessary requirement for making the right decision.

Decision Making↗

Differential proteomic analysis of Bacillus subtilis nitrate respiration and fermentation in defined medium.

A comparative investigation of protein expression by two-dimensional gel electrophoresis was conducted between Bacillus subtilis cultures grown in defined medium under aerobic, anaerobic nitrate respiration, or fermentation conditions. Defined medium specific for either nitrate respiration or fermentation allowed distinction between proteins induced by each individual growth process. Our differential protein profiling analysis between aerobic and anaerobic conditions showed that anaerobic fermentation induced at least 44 proteins and nitrate respiration induced at least 19 proteins compared to aerobic controls. Certain proteins were specifically induced during nitrate respiration or fermentation, while others were induced by both anaerobic processes. Eleven proteins induced by nitrate respiration and/or fermentation were identified by peptide mass matching using matrix-assisted laser desorption/ionization-time of flight mass spectrometry. Proteins encoded by feuA, hmp, and ytkD were induced by nitrate respiration. Proteins encoded by pyrR, sucD, trpC, and ywjH were induced by fermentation. Proteins encoded by acuB, pdhC, ydjL, and yvyD were induced by nitrate respiration and fermentation. This proteomic analysis has provided a more complete characterization of B. subtilis anaerobic growth and increased our understanding of its metabolic pathways of nitrate respiration and fermentation.

Bacillus subtilis↗

Respiratory protection: associated factors and effectiveness of respirator use among underground coal miners.

BACKGROUND: We investigated factors associated with the use of respiratory protection and explored the effectiveness of respirators among coal miners. METHODS: Between 1987 and 1992, respiratory symptoms, smoking, lung function, and dust exposures were assessed longitudinally among 185 underground bituminous coal miners. Self-reported use of respiratory protection was expressed as mean percent time wearing a respirator. RESULTS: Miners' respirator use increased with mean dust concentration, but decreased with tobacco consumption. Increasing age was associated with greater respirator use. Miners who had respiratory symptoms at the initial survey subsequently reported greater use of respirators. A significant protective association was found between the miners' respirator use and FEV(1) levels at both the initial and follow-up surveys. CONCLUSIONS: These results provide additional evidence that respirator use is protective of lung health. When respiratory protection programs are developed, factors that may affect respirator use behavior, such as age, smoking, and respiratory symptoms, should be considered. Future studies of respiratory health will need to consider workers' use of respiratory protection.

Adult↗

The carbon isotope composition of CO2 respired by trunks: comparison of four sampling methods.

The (13)C natural abundance of CO(2) respired by plants has been used in the laboratory to examine the discrimination processes that occur during respiration. Currently, field measurements are being expanded to interpret the respiration delta(13)C signature measured at ecosystem and global levels. In this context, forests are particularly important to consider as they represent 80% of the continental biomass. The objective of this investigation was to compare four methods of sampling the CO(2) respired by trunks for the determination of its carbon isotope composition: three in situ methods using chambers placed on the trunk, and one destructive method using cores of woody tissues. The in situ methods were based either on a Keeling plot approach applied at the tissue level or on an initial flush of the chamber with nitrogen or with CO(2)-free air. In parallel, we investigated the possibility of an apparent discrimination during tissue respiration by comparing the delta(13)C signature of the respired CO(2) and that of the organic matter. The study was performed on six tree species widely distributed in temperate and mediterranean areas. The four methods were not significantly different when overall means were considered. However, considering the individual data, the Keeling plot approach and the nitrogen flush methods gave fairly homogeneous results, whereas the CO(2)-free air method produced more variable results. The core method was not correlated with any of the chamber methods. Regardless of the methodology, the respired CO(2) generally was enriched in (13)C relative to the total organic matter. This apparent enrichment during respiration was variable, reaching as much as 3-5 per thousand. This study showed that, on the whole, the different sampling techniques gave similar results, but one should be aware of the variability associated with each method.

Carbon↗

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence.

We measured the response of dark respiration (R(d)) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communities in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R(d) at a reference temperature (R(10)) and the extent to which R(d) increased with temperature (described by E(o), a parameter related to the energy of activation, or the change in R(d) over a 10 degrees C range, Q(10)). Site averaged E(o) ranged from 44.4 kJ mol(-1) K(-1) at the 60-year-old site to 26.0 kJ mol(-1) K(-1) at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E(o) or Q(10)) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A(max)/R(d)) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.

Age Factors↗

Relationship between basal soil respiration rate, tree stand and soil characteristics in boreal forests.

Soil respiration is considered to represent the overall microbial activity reflecting mineralisation of organic matter in soil. It is the most commonly used biological variable in soil studies. In long-term monitoring of forested areas, there is a need for reference values for soil microbiological variables in different forest ecosystems. In this study we describe the relationship between soil respiration rate, tree stand and humus chemical characteristics of boreal coniferous forests stands. Soil respiration rate was higher in pine dominated than in spruce dominated study sites when the result was calculated on dry matter bases. However, when calculated on area bases, the result was opposite and no difference was found when the soil respiration rate was calculated on organic carbon bases. Irrespective of the main tree species, the soil respiration rate was equal in different development classes but not equal in soil fertility classes, i.e. within forest site types based on differences in ground vegetation. Respiration rates were clearly higher in mesic sites when calculated on dry matter, C(org) or area bases. However, soil respiration rate did not correlate with soil chemical variables indicating site fertility. Soil respiration rate on dry matter basis was at a lower level in the south and on more fertile sites, and on the other hand at a higher level in older stands and on sites with a thicker organic layer.

Carbon Dioxide↗