The role of microorganisms in mobilization and fixation of phosphorus in sediments

Cycling of phosphorus (P) at the sediment/water interface is generally considered to be an abiotic process. Sediment bacteria are assumed to play only an indirect role by accelerating the transfer of electron from electron donors to electron acceptors, thus providing the necessary conditions for redox-and pH-dependent, abiotic sorption/desorption or precipitation/dissolution reactions. Results summarized in this review suggest that

1. in eutrophic lakes, sediment bacteria contain as much P as settles with organic detritus during one year
2. in oligotrophic lakes, P incorporated in benthic bacterial biomass may exceed the yearly deposition of bioavailable P several times
3. storage and release of P by sediment bacteria are redox-dependent processes
4. an appreciable amount of P buried in the sediment is associated with the organic fraction
5. sediment bacteria not only regenerate PO₄, they also contribute to the production of refractory, organic P compounds, and
6. in oligotrophic lakes, a larger fraction of the P settled with organic detritus is converted to refractory organic compounds by benthic microorganisms than in eutrophic lakes.

From this we conclude that benthic bacteria do more than just mineralize organic P compounds. Especially in oligotrophic lakes, they also may regulate the flux of P across the sediment/water interface and contribute to its terminal burial by the production of refractory organic P compounds.     

Habitability of the early earth: Clues from the physiology of nitrogen fixation and photosynthesis

In the absence of direct evidence concerning the nature of the early Earth environments, it is acceptable under the uniformitarian principle to attempt to define primitive habitats from modern procaryotic physiology. Combining the rock and fossil record with present phylogenetic reconstuctions, application of this paleoecological approach to the evolutionary biochemistry and physiology of nitrogen fixation and photosynthesis leads to several inferences about the nature of Archean environments:

1. To stimulate nitrogenase evolution and avoid its repression, the activity of the NH ₄ ⁺ ion was less than 10^?3, and probably lower.
2. To be consistent with a moderately protective ozone screen, while not also repressing nitrogenase activity, incursions of abiotic dissolved oxygen at levels in the range 10^?1.2?10^?3.5 PAL would have been acceptable.
3. To induce the formation and activity of RuBP carboxylase, the pCO₂ was less than 100 PAL.
4. To support Photosystem I activity, sulfide concentrations of at least 10^?4 M were present in the photic zone.
5. To avoid a too-rapid oxidation of sulfide, the pH was probably between 6–7, where H₂S exceeds HS^?.

Evolutionary ‘pressure’ to stimulate the later development of oxygenic photosynthesis (Photosystem II), would require several subsequent habitat modifications:

1. Lowering the sulfide to < 10^?4 M to inhibit Photosystem I.
2. Raising the pH above neutral (HS^? > H₂S), to mediate more rapid oxidation of HS^?.
3. Maintaining either an illumination below 300–400 lux (to avoid photosynthetic O₂ self-repression of nitrogen fixation), or an adequate local source of combined nitrogen (aNH ₄ ⁺ > 10^?4) to repress nitrogen fixation entirely.

     

Effects of humic stress on the zooplankton from clear and DOC‐rich lakes

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Biogenesis of mitochondria 19 the effects of unsaturated fatty acid depletion on the lipid composition and energy metabolism of a fatty acid desaturase mutant of

1. The lipid composition of a mutant ofSaccharomyces cerevisiae which cannot synthesize unsaturated fatty acid (UFA) can be extensively manipulated by growing the organism in the presence of added fatty acids.
2. Growth of the mutant is supported by a wide range of unsaturated fatty acids including oleic, palmitoleic, petroselenic, 11-eicosaenoic, ricinoleic, arachidonic, clupanodonic, linoleic and linolenic acids; 9- and 10-hydroxystearic acids support growth less effectively, but erucic, nervonic, elaidic and saturated fatty acids (C_8∶0?C_20∶0)^* are ineffective. All the fatty acids which support growth are incorporated into cell lipids, apparently without further metabolism.
3. The effects of altered lipid composition on the energy metabolism of yeast cells were investigated. Cells containing less than approximately 20% of their fatty acids as UFA cannot grow on non-fermentable substrates, and their growth on glucose is restricted to that which can be supported by fermentation alone.
4. UFA-depleted cells contain mitochondria which are apparently normal in morphology, furthermore they have normal levels of cytochromesa+a ₃,b,c ₁ andc and respire at normal rates. This suggests that the lesion in energy metabolism produced by UFA-depletion may be the loss of the ability of the mitochondria to couple respiration to phosphorylation.
5. UFA-depleted cells incorporate added UFA into their cell lipids and subsequently regain the ability to grow on non-fermentable substrates, showing that the lesion in energy metabolism is fully reversible.

     

Characterization of solid and dissolved carbon in a spruce-fir Spodosol

Organic substances are an integral part of the biogeochemistry of many elements in forest ecosystems. However, our understanding of the composition, chemistry, and reactions of these materials are incomplete and sometimes inconsistent. Therefore, we examined in detail dissolved organic carbon (DOC) in forest floor leachates over a two-year period (1992–1993), soil C, and DOC adsorption by a mineral soil to determine the relationship between soil solid and solution C characteristics in a spruce-fir ecosystem. The structural composition of DOC, DOC fractions (hydrophobic and hydrophilic acids, hydrophilic neutrals), and soil samples from the organic and mineral horizons were also analyzed using¹³C nuclear magnetic resonance (NMR) spectroscopy.Total DOC in forest floor leachates ranged from 7.8 to 13.8 mmol L^–1 with an average of 8.6 mmol L^–1. Concentrations were highest in September of both 1992 and 1993. Fractionation of the forest floor DOC indicated these solutions contained high organic acid contents that averaged 92% of the total DOC. Hydrophobic acids were also preferentially adsorbed by the B horizon. The¹³C NMR data suggested alkyl, carbohydrate, aromatic, and carboxylic C were the primary constituents for organic and mineral soils, DOC, and DOC fractions. Compositional changes of C were observed as aromatic and carbohydrate decreased, whereas alkyl, methoxy, and carbonyl moieties increased with depth. However, C composition changed little among the three organic layers based on the similarity of alkyl/carbohydrates ratios as determined from NMR area integration, suggesting that in this acid soil, decomposition proceeds rather slowly. Hydrophobic acids contained high contents of aromatic C, whereas hydrophilic acids were comprised primarily of carboxylic C. Hydrophilic neutrals were rich in carbohydrate C. Results indicated that these DOC fractions were unaltered during the isolation process. Carboxylic C groups appeared to dissolve easily and were probably the primary contributor to organic acidity in our organic dominate leachates. Results also suggested that DOC materials adsorbed on the B horizon underwent further biodegradation. Several seasonal patterns of C composition were observed in the forest floor leachates and DOC fractions collected between 1992 and 1993.Overall, the evidence from this study suggested that (i) DOC levels were mainly controlled by biological activity, (ii) forest floor DOC was comprised primarily of organic acids, (iii) contact of soil leachates with B horizon material affected DOC quantitatively and qualitatively, (iv) phenolic, carboxylic, and carbonyl C appeared to dissolve readily in the forest Oa horizon, (v) DOC materials adsorbed on the B horizon selectively underwent further decomposition, and (vi) C composition is a function of the extent of decomposition and DOC fractions.     

Total recovery of O2 evolution and nanosecond reduction kinetics of chlorophyll-a II + (P-680+) after inhibition of water cleavage with acetate

Oxygen evolution and reduction kinetics of the photooxidized Chl-a_II ⁺ have been measured in oxygen-evolving complexes from the thermophilic cyanobacterium Synechococcus sp.

1. Incubation of PS II particles with acetate resulted in an inhibition of oxygen evolution and a retardation of the Chl-a_II ⁺=reduction kinetics from the nanosecond range to the microsecond range, indicating a modification of the donor side of photosystem II (PS II).
2. After the first two flashes given to a dark-adapted, acetate treated sample, Chl-a_II ⁺ was re-reduced with a half-life time of 160 μs by a component of the donor side of PS II. Under repetitive excitation Chl-a_II ⁺ was re-reduced in 500 μs by electron back reaction from the primary acceptor Q_A ^- (X-320^-). Obviously, in the presence of acetate only two electrons are available from the donor side.
3. Both oxygen evolution and nanosecond reduction kinetics of Chl-a_II ⁺ were restored to the control level when acetate was removed.
4. The results indicate a tight coupling between O₂ evolution and nanosecond reduction kinetics of Chl-a_II ⁺.
5. The reversible inhibition is probably due to a replacement of Cl^- by acetate within the water splitting enzyme.
6. Due to its strongly retarded kinetics, the reversibly modified system may facilitate investigations of the mechanism of the donor side.

     

Tracing carbon sources in small urbanising streams: catchment‐scale stormwater drainage overwhelms the effects of reach‐scale riparian vegetation

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Seasonal changes in ambient phosphate and phosphatase activities of the cyanobacterium Rivularia atra in intertidal pools at Tyne Sands, Scotland

1. A study was made of Rivularia atra at a sheltered bay in S-E. Scotland, where it is frequent in summer in pools of the upper eulittoral. Monthly measurements of water chemistry and phosphate activities were made at the same stage in the tidal cycle during April 1992–October 1993, together with other short intensive studies.
2. Drift seaweed deposited at high tide levels (>4.9 m) released high concentrations of inorganic nutrients to adjacent pools, where the water sometimes exceeded 6000 µg l^?1 PO₄?P. Combined inorganic N was, however, always low and the N:P ratio fell to 0.001–0.003:1 (by weight) in August, when the temperature was relatively high: much N was lost to the atmosphere as ammonia.
3. Total P concentrations in the Rivularia pools (mostly covered by 4.5–4.8 m tides) were much lower than in the drift seaweed pools, but similar to the sea, though with higher % organic P in the former (means of 50% v. 28%): total N concentrations were much lower in the Rivularia pools than in seawater, leading to a very low N:P for most of the time. Aqueous organic P tended to increase (in summer) during the period between tidal cover, even in pools lacking input drainage, suggesting that much of the organic P may be autochthonous.
4. All 9 Rivularia pools showed high concentrations of P in June in one or both years. At the same time the Rivularia colonies formed abundant hormogonia, followed 4–8 weeks later by a marked increase in macroscopically obvious colonies.
5. Assays of ‘surface’ phosphomonoesterase activity by Rivularia colonies showed different results according to whether p-nitrophenyl phosphate (pNPP) or 4-methylumbelliferyl phosphate (4-MUP) was used as substrate. Both substrates indicated obvious activity at intervals during the year. Using pNPP, activity was very low in June, usually followed by a very high value in July or August; use of 4-MUP showed a different response, with a high value one or two months later than with pNPP.

     

Changes in the chemical composition of alder,poplar and willow leaves during decomposition in a river

1. As there was a paucity of information on the aquatic decomposition of leaves of willow (Salix alba), poplar (Populus gr. nigra) and alder (Alnus glutinosa), the net-bag technique was used to study chemical changes in the leaves over a period of six months in the Garonne (France).
2. The disappearance rates of foliar organic matter were about similar for the three species (k = 0.0065 day^?1 for alder, k = 0.0054 day^?1 for poplar and k = 0.0050 day^?1 for willow). Changes in carbon amount were comparable to those of organic matter. Nitrogen and organic phosphorus accumulated during the first months of decomposition. The C/N ratios of foliar matter changed in the same way in the three species, falling steeply for the first month, then levelling off. Amounts of sugar disappeared very rapidly, particularly for alder and willow. Cellulose concentrations were constant throughout the whole decomposition process whilst an increase in the amounts of lignin was observed during the first months; however the latter could be due to an interference by microbial compounds.
3. The changes of some constituents, such as carbon, sugars and cellulose, were described with exponential or multiexponential models. Despite the very different initial ratios of C/N and lignin/N, these changes associated with leaf decomposition were similar for the three species.

     

Role of silicon in diatom metabolism

1. In silicic acid-starved cells of the diatom Nitzschia alba, ⁶⁸Ge(OH)₄ is transported against a concentration gradient, leading to intracellular concentrations of germanic acid up to 3500 times greater than the exogenous concentrations. The accumulated substrate is osmotically active, as determined by its efflux into germanic acid-free medium.
2. Metabolic energy is required for Ge(OH)₄ transport, since uptake is completely inhibited by 1 mM DNP, 5×10^-2 M sodium azide or 1 mM iodacetamide, and is strongly inhibited by CCCP and antimycin A. Inhibition of protein synthesis with 20 μg/ml cycloheximide does not affect the initial velocity of transport, but strongly reduces the steady state intracellular concentration.
3. A double reciprocal plot of uptake velocity versus substrate concentration yields a biphasic curve. The kinetic data are consistent with the interpretation that N. alba has two transport systems for germanic acid; a high affinity-low capacity (K _s=0.36 μM; V _max 1.2 μmoles/10⁸ cells/min) system and a low affinity-high capacity (K _s=5 μM; V _max 6.2 μmoles/10⁸ cells/min) system.
4. The implications of these findings for silicic acid transport and metabolism in N. alba are discussed.

     

An ecological study of the Ponggol Estuary in Singapore

1. An ecological study of the Ponggol Estuary was conducted from July 1965 to June 1966 and the seasonal data on physical, chemical and biological characteristics were presented.
2. The Ponggol River represents a short, narrow and shallow estuary in Singapore. The river mouth is open throughout the year and water from eastern Johore Straits drains in twice a day at high tide. The upper reach, however, is left exposed at low tide.
3. The Ponggol River was classified as a vertically and laterally homogeneous estuary and was found to exhibit a mesohaline to polyhaline environment.
4. Significant hydrological gradients from the river mouth 10 the upper reach were noted in the river system. Salinity, dissolved oxygen and pH increased towards the mouth of the river and other parameters such as nutrients, dissolved organic matter and turbidity increased towards the source.
5. Although the river received organic pollutants at the upper reach the estuary was able to discharge them fairly rapidly through regular flushing by the tides. The transient rise of organic matter did not appear to impart any serious affect on the biota in the estuary.
6. Over 98% of the phytoplankton consisted of diatoms, most of which were brought into the estuary from eastern Johore Straits. Freshwater forms were relatively few.
7. Phytoplankton biomass was considerably higher than the adjoining waters. and was reduced at the upper reach due to high turbidity of the water.
8. 80% of the zooplankton was composed of dinoflagellates,Difflugia, copepods and bivalve larvae dominating at all sections of the estuary.
9. Percentage composition of the zooplankton showed that dinoflagellates and copepod nauplii predominated at high tide whileDifflugia and bivalve larvae were abundant at low tide.
10. Zooplankton standing crop, in general, was higher towards the source at high tide but the reverse was found at low tide, i.e. standing crop increased towards the river mouth. This was attributed to the process of concentration.
11. Species composition of zooplankton was found to be more or less similar to that of the eastern Johore Straits.
12. The nekton consisted predominantly of small and juvenile fish. Close correlation of fish and copepods was found to be statistically valid and it was concluded that the fish entered the estuary to feed rather than to spawn.
13. The squids formed an important catch of the beach seine unit and were caught throughout the year.
14. The fish population could be grouped into four categories: estuarine components, euryhaline components, marine components and migratory components.
15. Benthic invertebrates were abundant. Commercially important species consisted of prawns,Metapenaeus andPenaeus, and crabs,Neptunus pelagicus andScylla serrata.
16. The river bed was inhabited predominantly by molluscs and the distribution resembled that of the sheltered shore of muddy-sand type.

     

Degradability, molecular weight and adsorption properties of dissolved organic carbon and nitrogen leached from different types of decomposing litter

Aims

We characterized dissolved organic matter (DOM) leached during decomposition of deciduous silver birch litter (Betula pendula Roth.), coniferous Norway spruce litter (Picea abies (L.) Karst.) and a mixture of these litters in order to find out whether the properties of DOM would explain the earlier observed signs for higher microbial activity in soil under birch than spruce.

Methods

DOM leached from decomposing litters was collected in a litter-column experiment in the laboratory. Adsorption properties (XAD-8 resin fractionation) and molecular weight as well as the degradability of dissolved organic carbon (DOC) and nitrogen (DON) were measured three times during decomposition: 1) in the early stages, 2) after the mass loss reached 20–30 % and 3) when the mass loss reached 30–40 %.

Results

The leaching of DOC hydrophilic neutrals and bases, regarded easily degradable, decreased during decomposition. The leaching of DOC in hydrophobic acids, regarded refractory, increased from spruce and especially from the mixture litter during decomposition and may be connected to the degree of litter decomposition that was highest for the mixture. Unexpectedly, the degradability of DOC differed only slightly between the litters but the degradability of DON was substantially higher for spruce than birch. Spruce DOM seemed to be more N-rich than birch DOM in the early stages of decomposition and it seemed that labile DON was mobilized earlier from spruce than birch litter.

Conclusions

We conclude that the decomposition degree of litter determines largely the properties of DOM. The observed differences in the properties of DOM sampled during the litter decomposition cannot explain differences in C and N cycling between birch and spruce.     

Muscle GSH-Px activity after prolonged exercise,training, and selenium supplementation

A double-blind study of the effects of supplementing with selenium vs. placebo on the physiological responses to acute and chronic exercise was conducted in 24 healthy, nonsmoking males, mean age 22.9±2.1 yr, randomly divided into two groups of 12 (Pla/Sel). After a controlled period in the absence of training, all subjects were put on an individualized endurance training program with the same rules of progression and overload (3 sessions/wk×10 wk). Supplementation, either real (240 μg of organic selenium/d in Sel group) or imaginary (Pla group) was administered during the same period. In each of the conditions Pre- and Post- (training ± sel supplementation), muscle, plasma, and systemic parameters were determined before (BF) and after (AF) acute exercise, involving the repetition of muscle work cycles separated by 5-min recovery periods, combining 20 min at 65% and a maximal duration of 100% VO₂ max of running on a treadmill, leading the subjects to exhaustion between 2 h 40 min and 3 h 30 min. Changes in parameters as a function of three independent variables:

1. Acute exercise (E);
2. Chronic exercise (T); and
3. Selenium supplementing (S)

were tested with ANOVA and the Student\rsst-test on paired series. Among the variables examined, muscle glutathione peroxidase (GPx) presented a remarkable behavior. Enzymatic activity:

1. Decreased significantly (p<0.05,n=24) between the beginning and the end of acute exercise: 29.6±12 vs. 20.8±8.1 IU·g protein^?1 in Pre conditions;
2. Remained at the same level in resting conditions between the beginning and end of training (from Pre to Post) regardless of the group: 33.5±10.8 vs. 32.3±19.8 and 25.7±12.4 vs. 23.5±10.2 IU·g protein^?1 in Pla and Sel subjects, respectively; and
3. Increased from 23.5±10.2 to 37.3±28.5 (P=0.057) during acute exercise in Post-conditions (after training) in supplemented subjects (Sel group).

The situation was as if acute exercise played the role of allosteric stimulator of the GPx reaction in muscle.     

The role of touch,pressure and nociceptive mechanoreceptors of the leech in unrestrained behaviour

1. The maximum force exerted against an isometric force transducer by 6 leeches weighing 2.6–3.7 g, as they squeezed through apertures of different widths varied inversely with aperture width.
2. T cells in the leech skin code for velocity of indentation, not pressure or displacement. The frequency with which T cells fire is best described by two log functions, one for low, another for fast indentations. T cells responded to indentation velocities down to 10 μms^?1.
3. The average threshold pressure for 5 P cells was 150 kPa and for 5 N cells was 521 kPa.
4. We conclude from these data that when leeches explore their mechanical environment and initiate contact with external objects, the threshold pressure for N cells is rarely crossed. Of the three classes of mechanoreceptor, T cells are the main modality through which leeches obtain contact information, though P cells may occasionally be recruited for local pressure peaks.

     

Floristic composition and phenology of temperate grasslands of Western Himalaya as affected by scraping,fire and heavy grazing

This paper reports the analysis of floristic composition and phenology of protected, annually scraped and fired, and heavily grazed grasslands community of Western Himalaya.

1. A total of 19, 20 and 23 species were recorded at protected (S₁), annually scraped and fired (S₂) and heavily grazed (S₃) sites respectively.
2. Domination of therophytic flora as indicated by biological spectrum study, was noted in all the three sites.
3. Density of plants reached a maximum in the rainy season and amounted to be 4300.5, 3439.8 and 3298.0 tillers/m² on S₁, S₂ and S₃ respectively.
4. Maximum total basal cover was also recorded in the rainy season and amounted to be 769.9, 3390.6 and 2295.7 cm²/m² on S₁, S₂ and S₃ respectively.
5. Correlation between density and basal cover was positive on all three sites.

     

Effects of in vivo ethanol administration on Ca2+/Mg2+ ATPase and ATP-dependent Ca2+ uptake activity in synaptosomal membranes

1. Acute administration of ethanol (4 g/kg, i.p.) to mice inhibits the sequestration of calcium into endoplasmic reticulum-like organelles in synaptosomal membranes.
2. Ethanol administration inhibits both Ca²⁺-stimulated adenosine triphosphate hydrolysis and ATP-dependent calcium uptake in the vesicles at time of loss of righting reflex.
3. At recovery of righting reflex, the Ca²⁺-ATPase activity returns to normal levels, while the ATP-dependent uptake remains inhibited.
4. The effect of ethanol is specific for the sequestration (active transport) of calcium since calcium binding to synaptic membranes is not altered.
5. Alteration in mechanisms responsible for synaptosomal buffering of cytosolic Ca²⁺ levels by in vivo ethanol may contribute to altered transmitter release rates following ethanol adminstration.

     

Limnology and trophic status of glacial lakes in the tropical Andes (Cajas National Park,Ecuador)

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Seasonal variations in biochemical constituents of the clam,Paphia laterisulca

1. The total protein, fat and glycogen contents were estimated from the edible clam, P. laterisulca. Seasonal variations in these constituents along with the water content were studied.
2. The gonad index in P. laterisulca was found to increase during the ripe condition and in winter (December–January) and decrease on spawning.
3. A relatively high water content was obtained during monsoon (June to September). This might be due to the loss of salts and gain of water in low salinities.
4. Protein content varied with the reproductive cycle of the clam. The level reached its peak in the mature stage and declined on spawning. Immature clams showed less protein content than gravid ones.
5. Lipid content started to increase as the gametogenesis commenced, reached its peak in fully mature condition (August) and sharply declined due to the shedding of gametes during spawning.
6. Glycogen content was high during the period of active gameto-genesis (May–June). A sharp decrease took place when the clams were fully ripe (July). The glycogen might have been utilized in the formation of active ripe gametes.
7. After starvation for twelve days, total protein and fat contents remained constant, while glycogen content decreased by 66.82%. The water content increased by 4.67%.
8. Seasonal variation in the organic constituents are discussed in relation to the reproductive cycle of the clam.

     

Acid-base titration across the plasma membrane ofMicrococcus denitrificans: Factors affecting the effective proton conductance and the respiratory rate

The object of this work was to measure the effective proton conductance of the plasma membrane ofMicrococcus denitrificans under various conditions and to investigate possible connections between respiration and proton translocation.

1. Pulsed acid-base titrations of suspensions ofM. denitrificans in a medium containing the permeant thiocyanate ion, or when K⁺ ion permeability was induced by valinomycin in a KCl medium, showed that the normal effective proton conductance of the membrane system was less than 1 μmho/cm².
2. A pH-overshoot artefact was suppressed by adding carbonic anhydrase.
3. The effective proton conductance was increased by the uncoupler FCCP in the same concentration range as was required to stimulate respiration. Concentrations of FCCP above 1·5 μM inhibited respiration after an initial stimulation.
4. The effective proton conductance in presence of 2 μM FCCP was at least 17 μmho/cm².
5. The quantitative relationships between the respiratory rate, the stoichiometry of respiration-driven proton translocation, and the effective proton conductance of the membrane of the cells are compatible with the suggestion that stimulation of respiration by FCCP is due to a release of back-pressure exerted by a protonmotive potential on the respiratory chain system in the membrane. Only one amongst other possible explanations of the stimulation of respiration by FCCP is, however, excluded.

     

Respiration-driven proton translocation in micrococcus denitrificans

The polarity and stoichiometry of respiration-driven proton translocation was studied by electrometric and spectrophotometric techniques inMicrococcus denitrificans in the context of the energy transduction mechanism in bacterial oxidative phosphorylation.

1. Protons are ejected through the plasma membrane during respiratory pulses and thereafter diffuse slowly back.
2. In presence of ionic species mobile across the membrane (K⁺-valinomycin, K⁺-gramicidin, or SCN^?), limiting→H⁺/O quotients of 8 were obtained with endogenous respiratory substrates, and the rate of translocation (14·3 μg ions of H⁺/sec g cell dry weight) was commensurate with that of respiration optimally stimulated by FCCP at an →H⁺/O quotient of 8.
3. The rate of decay of the proton pulses was greatly increased by FCCP, but there was little or no effect on the →H⁺/O quotient characteristic of the respiratory system.
4. Various interpretations of the observations are discussed, and it is concluded that respiration is probably coupled directly or indirectly to electrogenic proton translocation. The observations are compatible with the chemiosmotic hypothesis of coupling between respiration and phosphorylation.