Control of acid-base status in active and dormant land snails,Otala lactea (Pulmonata,Helicidae)

Summary Control of extracellular acid-base status was examined during activity and dormancy inOtala lactea (Pulmonata, Helicidae). Active snails showed little variation in hemolymph pH and at constant temperature. With increase of temperature, hemolymph increased from about 6 Torr at 5°C to 13 Torr at 24°C and pH decreased by about 0.017 pH units/°C, a pattern consistent with alphastat regulation of pH via ventilatory control of .During dormancy, mean hemolymph increased to about 50 Torr. Venous pH declined by about 0.4 units due to hypercapnia and fluctuated more widely than in active snails due to variability of . Hemolymph pH declined further in prolonged dormancy due to progressive metabolic acidosis; after one year of dormancy the mean hemolymph pH was about 0.8 units lower than that of active snails at similar temperature.Active snails exposed experimentally to high showed a large increase in hemolymph [HCO ₃ ^– ]. However, [HCO ₃ ^– ] declined by up to 50% during dormancy, despite the naturally occurring hypercapnia. Hemolymph osmolality and the concentrations of solutes other than [HCO ₃ ^– ] increased with increasing duration of dormancy. Concentrations of magnesium and calcium increased about 2.5 times more rapidly than those of sodium and chloride, indicating that acidosis is partially offset by the dissolution of carbonates from the shell or tissues.     

Growth and product formation of actinomycetes cultivated at increased total pressure and oxygen partial pressure

Summary Influence of pressure (P) and oxygen partial pressure ( ) on cultivation of various Streptomyces spp. and Micromonospora purpurea was examined in pressurized air-lift and stirred tank fermenters. The maximum was 2100 mbar. Growth and product formation of all cultures tested were markedly influenced by higher than 1000 mbar. There is evidence that wild strains are more oxygen tolerant than production strains. At a certain the metabolic activities of all cultures were inhibited. However, results obtained with S. aureofaciens and S. rimosus indicated an increase in specific product formation rate at elevated pressure. With increase in oxygen tension incorporation of oxygen into tetracycline molecules was enhanced. Since elevated oxygen tension can either show inhibiting effects or may be used for regulation of product formation and selectivity, the influence of should be determined in an appropriate experimental set-up for each process.Offprint requests to: U. Onken     

Changes in the respiratory properties of the blood in the carp,Cyprinus carpio, induced by diurnal variation in ambient oxygen tension

Summary Effects of diurnal variation in ambient oxygen tension on acid-base balance and blood respiratory properties were investigated in carp (Cyprinus carpio). The carp were subjected to two cycles in ambient between about 130 mm Hg and about 12 mm Hg at 17°C (cf. Figs. 1 and 2). The first period of hypoxia was characterized by a non-compensated respiratory alkalosis, i.e. whole blood showed an increase in pH from 7.92 to 8.14. During the second hypoxic period, 24 h later, a significantly smaller respiratory alkalosis was present, whole blood pH changed from 7.95 (the value found during the intermediate return to normoxia) to 8.08. The latter increase was associated with a significant increase of 25% in plasma bicarbonate concentration compared with the first period of hypoxia (Fig. 1, Table 1). The erythrocytic concentrations of hemoglobin and ATP were lowered by about 10%, compared with the normoxic values, during the two episodes of hypoxia, and this was due to a swelling of the erythrocytes during hypoxia. The red cell GTP concentration showed an altogether different change during the O₂ cycling: the absolute concentration of red cell GTP changed to a steady level, 50% below that present at the onset of the experiment and the major part of this change took place between the two hypoxic periods (Fig. 2, Table 2).The results are discussed with reference to the respiratory function of the blood of carps during subjection to cyclic changes in nature.     

Activity and oxygen consumption during hypoxic exposure in high altitude and lowland sceloporine lizards

Summary Resting rates of O₂ consumption against , exercise endurance times and during recovery from vigorous exercise were measured inSceloporus occidentalis captured near sea level and inS. graciosus captured above 2850 m. Oxygen consumption against was also measured inS. occidentalis captured above 2850 m. When was recorded continuously, as ambient was slowly reduced from 155 Torr, it became directly dependent upon ambient between 110 and 120 Torr. The critical for the high altitude lizards was lower than that for the lowland lizards, which enabled the former to maintain relatively higher 's when ambient was reduced below 120 Torr. The high altitude lizards also had significantly greater endurance when stimulated to exercise at 1600 m ( 130 Torr). Both the higher under hypoxia and the greater endurance roughly parallel a significantly greater maximum in the high altitude lizards. At a simulated altitude of 3600 m ( 100 Torr), maximum and rate of recovery of the O₂ debt calculated from post active were significantly reduced in the lowland but not the high altitude lizards. The effects of simulated altitude conditions on the lowland but not the mountaine animals indicate adaptations to altitude in these sceloporine lizards. We did not find any consistent relationship between organ/body weight ratios or hematocrit and our measures of endurance or the altitude at which the lizards were captured.     

Reversed bohr and root shifts in hemocyanin of the marine prosobranch,Buccinum undatum: Adaptations to a periodically hypoxic habitat

Summary Oxygen binding properties of the hemocyanin-containing blood ofBuccinum undatum were examined in vitro and in vivo under normoxic ( 150 mmHg) and hypoxic ( 50 mmHg) conditions at 10°C. Blood pH and showed a decrease in vivo under hypoxic conditions. Oxygen uptake at high water , was about 18 ml O₂/kg·h (wet weight) and the critical oxygen tension between 25 and 50 mm Hg. In vitro the O₂ binding to hemocyanin showedn-values independent of pH, while both O₂ affinity and oxygen carrying capacity were strongly pH dependent. Oxygen affinity increased below pH=8.1 and thus showed a pronounced reversed Bohr shift in the physiological pH range (7.5<pH<8.1). The oxygen carrying capacity similarly increased markedly with falling pH in the physiological pH range (reversed Root shift). Astrup titration curves showed a metabolic and respiratory acidosis under hypoxic conditions ( 50 mm Hg). The role of hemocyanin in the transport of oxygen in relation to ambient O₂ availability is discussed.     

Gas exchange in the incubation mounds of megapode birds

Summary The brush turkey (Alectura lathami) and mallee fowl (Leipoa ocellata) are megapode birds that incubate their eggs by burying them in mounds. Respiratory gas exchange between the buried eggs and the atmosphere occurs mainly by diffusion through about 60 cm of decomposing forest litter (brush turkey) or sand (mallee fowl).Gas fluxes in the brush turkey mound are greatly influenced by the respiration of thermophilic microorganisms which consume O₂ at rates over eight times that of all of the eggs. The respiratory exchange ratio ( ) of the microorganisms is 0.75 and theQ ₁₀ for metabolism is 2.56. Fermentation and nitrogen fixation do not occur in the mounds.If the mound becomes too wet, gas tensions near the eggs can become critical because water increases rates of microbial respiration and impedes gas diffusion. However, field mounds are relatively dry, possibly because the adult bird modifies the shape of the mound and affects the entry of rain water. At egg level in field mounds, and are about 132 and 21 Torr, respectively, in both species. Embryonic respiration decreases and increases about 5 Torr in the immediate environment of individual eggs in late development. Due to a high eggshell gas conductance, which increases during incubation, the gas tensions within the shell of late embryos ( ca. 108 Torr, ca. 47 Torr) are not far from the mean values found in species that nest above ground.     

Water economy in bird nests

Summary Nest humidity ( ) under an incubating bantam hen was measured at ambient ranging from 1.3 to 25.0 mmHg. Weight loss of eggs was recorded in clutches of varying size. Nest and ambient were also measured in wild incubating willow ptarmigan nests in dry and wet habitats.Nest increased linearly with ambient in a way predictable on the assumption that the water vapour conductance ( ) of brood patch skin, plumage and eggs were constant and independent of ambient . Nest was also dependent of clutch size. Egg dehydration was quantitatively predicted from measured values and the laws of diffusion.Our findings confirm earlier conclusions that the adult bird does not actively regulate nest at varying ambient . Birds can presumably achieve appropriate egg dehydration by a strategy combining nest site, nest construction, egg shell conductance and incubation behaviour which meets the requirements of their breeding climate.Abbreviations water vapur pressure - water vapur conductance - water flux     

Respiratory responses to long-term temperature exposure in the box turtle,Terrapene ornata

Summary In late February, seven box turtles were collected with body temperatures between 7 and 9°C. Ventilation, gas exchange and end-tidal and were recorded at 5, 10, 15 and 25°C, the animals being at each temperature for 2 to 3 weeks. There was a pronounced diurnal rhythm of breathing frequency at all temperatures. At 5°C the mean 24-h frequency was only 3.7 breaths h^–1. At 15°C the frequency was 16 times higher with a 17-fold increase of ventilation. Oxygen uptake only changed from 3.4 to 12.7 ml·kg^–1·h^–1. Consequently, the ratio (ventilation, ml BTPS/O₂ uptake, ml STPD) increased from 12.5 at 5°C to 48 at 15°C, but decreased to 24 at 25°C. The decrease of this ratio during cold exposure contrasts with an increase of the ratio during cooling earlier reported for fresh water turtles,Pseudemys. Cutaneous CO₂ elimination was important at low temperature. This caused a decrease of the pulmonary gas exchange ratio so that end-tidal remained low at 5°C in spite of an end-tidal of only 54 Torr.     

A simple method to quantitatively measure polypeptide JHNHα coupling constants from TOCSY or NOESY spectra

A simple linear relationship between the J coupling constant and the linewidth (_1/2) of in-phase NMR peaks has been identified. This relationship permits the rapid and accurate determination of polypeptide J coupling constants from a simple inspection of amide cross peaks in homonuclear ¹H TOCSY or ¹H NOESY spectra. By using the appropriate set of processing parameters we show that J = 0.5(_1/2) – MW/5000 + 1.8 for TOCSY spectra and J = 0.6(_1/2) – MW/5000 – 0.9 for NOESY spectra, where _1/2 is the half-height linewidth in Hz and MW is the molecular weight of the protein in Da. The simplicity of this relationship, combined with the ease with which _1/2 measurements can be made, means that J coupling constants can now be rapidly determined (up to 100 measurements in less than 30 min) without the need for any complex curve-fitting algorithms. Tests on 11 different polypeptides involving more than 650 separate J measurements have shown that this method yields coupling constants with an rmsd error (relative to X-ray data) of less than 0.9 Hz. Furthermore, the correlation coefficient between the predicted NMR coupling constants and those derived from high-resolution X-ray crystal structures is typically better than 0.89. These simple linear relationships have been found to be valid for peptides as small as 1 kDa to proteins as large as 20 kDa. Despite the method's simplicity, these results are comparable to the accuracy and precision of the best techniques published to date.     

Electrochemical potential of protons in vesicles reconstituted from purified,proton-translocating adenosine triphosphatase

Summary Measurements were made of the difference in the electrochemical potential of protons ( ) across the membrane of vesicles reconstituted from the ATPase complex (TF ₀ ·F ₁) purified from a thermophilic bacterium and P-lipids. Two fluorescent dyes, anilinonaphthalene sulfonate (ANS) and 9-aminoacridine (9AA) were used as probes for measuring the membrane potential () and pH difference across the membrane ( pH), respectively.In the presence of Tris buffer the maximal and no pH were produced, while in the presence of the permeant anion NO ₃ ^– the maximal pH and a low were produced by the addition of ATP. When the ATP concentration was 0.24mm, the was 140–150 mV (positive inside) in Tris buffer, and the pH was 2.9–3.5 units (acidic inside) in the presence of NO ₃ ^– . Addition of a saturating amount of ATP produced somewhat larger and pH values, and the attained was about 310 mV.By trapping pH indicators in the vesicles during their reconstitution it was found that the pH inside the vesicles was pH 4–5 during ATP hydrolysis.The effects of energy transfer inhibitors, uncouplers, ionophores, and permeant anions on these vesicles were studied.     

Characterization of Four Novel Variants of Goat βA-Globin Gene

Four novel alleles of the adult -globin gene of Capra hircus were observed in an extended study on hemoglobin polymorphism in goat breeds living in the island of Sardinia. Nucleotide sequencing showed that one of these alleles is due to a 2 bp substitution at codon 125 ( G, "LeuGlu). Two substitutions, the silent CT for Leu at codon 78 and the conservative A G (Lys Arg) at codon 104, are shared by the other three alleles, two of them having additional mutations, which suggests a common origin. The allele we provisionally called the ^Y shares four out of five amino acid substitutions, together with the same polymorphisms in the IVSII, we observed previously in the rather common ^E gene. This evidence allowed the origin of the ^E gene to be better characterized. The data increase to seven the number of alleles at the goat ^A -globin locus characterized thus far at the molecular level. A simplified nomenclature for the increasing number of goat -globin alleles is presented.     

Ventilation and gas exchange in the diamondback water snake,Natrix rhombifera

Summary Simultaneous measurements of pulmonary and cutaneous oxygen and carbon dioxide exchange, pulmonary ventilation and heart rate were made on the diamondback water snake,Natrix rhombifera at 28°C using body plethysmography. Resting lung volume, maximum lung volume and tracheal volume were also measured.The following mean values were measured in undisturbed snakes breathing room air: total (pulmonary and cutaneous) O₂ uptake 46 mol · (kg min)^–1; total CO₂ output, 49 mol · (kg min)^–1; tidal volume, 12 ml (BTPS) · kg^–1; ventilatory rate, 6.9 min^–1; heart rate, 42 min^–1. From the measurements of tracheal volume, the effective (alveolar) ventilation was estimated as approximately 70% of total ventilation resulting in effective pulmonary and of 130 Torr and 20 Torr respectively. Cutaneous exchange accounted for 8.1% of the total and 12.4% of the total .Resting lung volume of anaesthetized snakes was 75 ml (BTPS) · kg^–1, maximum lung volume was 341 ml (BTPS) · kg^–1 and tracheal volume was 3.9 ml (BTPS) · kg^–1.     

Influences of exercise-stress and adrenaline upon intra- and extracellular acid-base status,electrolyte composition and respiratory properties of blood in tench (Tinca tinca) at different seasons

Summary Exercise-stress in tench resulted in severe acidoses in both the red cells and the extracellular fluid in vivo. These coincident pH decreases conformed to the in vitro pH_i-pH_e relationship for tench blood in the oxygenated state. The extracellular acidosis was primarily respiratory in winter and metabolic in spring and summer. This was due to more effective buffering of metabolic protons in winter by an elevation in [HCO ₃ ^– ] levels, rather than to differences in the lactic- and carbonic acid loads. A good correspondence was found between buffered metabolic protons and increases in [lactate].There was no evidence for -adrenergic red cell swelling and associated red cell pH changes in tench both after exercise and adrenaline infusion. Arterial O₂ transport was, however, improved in exercise by pronounced increases in .Large increases in plasma potassium concentration and small elevations of chloride and calcium levels occurred in exercise. Hematocrit and blood [Hb] also increased, probably due to an adrenergic release of erythrocytes from the spleen, but these increases were small and appeared unimportant for blood O₂ transport.Seasonal differences were found in exercise-induced changes in [lactate], in the magnitude of electrolyte and changes, as well as in resting values for pH_e, pH_i, [HCO ₃ ^– ], [Cl^–] and [Ca⁺⁺]. The origin and importance of these are discussed.     

Investigation of the structure of two-phase flow model-media in bubble-column bioreactors

Summary By applying photographic, electrical conductivity, and electrooptical methods, the transverse variation of bubble size and velocity, the local gas hold up, and the local specific gas/liquid interfacial area were estimated in a bench scale bubble-column bioreactor containing model cultivation media. The liquid velocity profile, the transverse turbulence intensity variations, and the turbulence energy dissipation scale were also measured by a hot film turbulence probe and constant temperature anemometer technique.A significant relationship was found between the two-phase flow fluid dynamical state and the transverse variation of the various properties.Symbols M mass - L length - T time - a gas/liquid interfacial area L² - specific gas/liquid interfacial area with regard to the bubbling layer volume L^–1 - D transverse coordinate (measured from the wall of the column) L - d bubble diameter L - d mean bubble diameter L - d_e dynamic equilibrium (maximum stable) bubble diameter L - d_p primary bubble diameter L - d_s Sauter bubble diameter L - E specific energy dissipation rate with regard to the volume of the liquid ML^–1T^–3 - EV_L energy dissipation rate ML²T^–3 - , since =1 g/cm³, E has the same numerical value as E. Therefore, the symbol E is used everywhere in the present paper for E for simplicity and called energy dissipation rate (S.s^–2=Stokes.s^–2) L²T^–3 - E_G or local relative gas holdup - f (r) cross correlation function - g acceleration of gravity LT^–2 - h longitudinal distance from the aerator L - relative turbulence intensity - N_O number of u and crossings T^–1 - n_B bubble frequency T^–1 - r distance between two points 1 and 2 of the cross correlation function L - t time - u instantaneous liquid velocity LT^–1 - mean liquid velocity LT^–1 - mean square fluctuation velocity L²T^–2 - turbulence intensity LT^–1 - w_SG superficial gas velocity LT^–1 - w_SL superficial liquid velocity LT^–1 - or E_G local relative gas holdup LT^–1 - energy dissipation scale L - kinematic liquid viscosity L²T^–1 - liquid density M L^–3 - surface tension M T^–2 - dynamic turbulence pressure M L^–1T^–2 Indices p primary (at the aerator) - e equilibrium (far from the aerator)     

Differentiation between acetate and higher volatile acids in the modeling of the anaerobic biomethanation process

This paper presents a model for the single-stage completely-mixed anaerobic digestion of complex substrates containing no volatile acids. In the model, volatile acids produced by the acidogenic bacteria are no longer considered together. Acetate is assumed to be representative of the substrate and propionate and butyrate act only as inhibitors for the methanogenic bacteria.Nomenclature ···0 represents factors associated with the influent - ···1 represents factors associated with the acidogenic bacteria - ···2 represents factors associated with the methanogenic bacteria - Q hydraulic flow (1/d) - V reactor liquid volume (1) - T temperature of the mixed liquor (° C) - S microorganisms concentration (mg/1) - L volatile solids concentration (mg VS/1) - Lb biodegradable volatile solids concentration (mg VS/1) - VA2 acetate concentration (mg/1) - VA3 volatile acids with 3 to 5 carbon atoms concentration (mg/1) - methane rate production (1 CH₄/1_digester.d) - Km saturation coefficient (mg/l) - Ki inhibition coefficient (mg/l) - specific growth rate (1/d) - maximum specific growth rate (1/d) - b biological decay coefficient (1/d) - Arrhénius coefficient (–) - yield of acidogenic bacteria per mg of biodegradable matter consumed (mg S₁/mg L_b) | (mg S₂/mg VA2) - yield of methanogenic bacteria per mg of VA2 consumed - yield of methane production per mg of S₂ formed (1 CH₄/mg S₂) - proportion of VA2 produced per mg of S₁ biosynthetised - proportion of VA3 produced per mg of S₁ biosynthetised - Y_sp volume of CH₄ produced per g of volatile solids eliminated     

Effects of ambient and local cutaneous temperatures on hand blood flow

The hand blood flow ( ) was investigated in response to a wide range of general and local cutaneous thermal stimuli (0–36°C and 4–42°C respectively), the local stimulus consisting of a thermostatically controlled water bath for the right hand (T_w), and the general stimulus, the ambient room temperature (T_a). was measured at the right wrist by strain gauge plethysmography; it was seen to respond more significantly to variations in T_w than to those in T_a at cold to comfortable ambient temperatures (T_a<22°C). A paradoxical vasodilatation was observed at T_w=4°C (Lewis' hunting phenomenon). The graphs of versus T at average to high local cutaneous temperatures (T_w > 33°C) are remarkably similar, except for an upward shift at successively higher values of T_w. The slope (or vasomotor reactivity) is interpreted as being controlled by variations in T_a. The curves exhibited maximum values at T_a = 31°C. Their subsequent decrease could represent a thermoregulatory adaptation to environment-organism heat transfer, the relative vasoconstriction tending to reduce the transfer. Although the qualitative response was the same for both sexes, the absolute value of was generally greater in male than in female subjects.     

Sodium influx in isolated gills of the freshwater mussel,Ligumia subrostrata

Summary Isolated gills of the freshwater mussel,Ligumia subrostrata, accumulate Na from a pondwater bathing medium. The rate of Na transport by the isolated gill is 13.2±1.1 mol (g dry gill·10 min)^–1 which equals or exceeds the estimated Na transport rate of intact animals. Sodium influx is saturable with aV _max of 13.6±1.2 mol (g dry gill·10 min)^–1 and an affinity (K _s) of 0.17 mM Na/l. The isolated gills survive prolonged exposure to pondwater with a constant of 890 l O₂ (g dry gill·h)^–1 over a 4 h period. Sodium transport in the isolated gills is stimulated 80% above control values by 10^–4 M serotonin, 60% by 0.5 mM cAMP and 60% by 12.5 g/ml nystatin. Sodium influx is inhibited by 0.5 mM amiloride and 1 mM lithium.     

Induction of somatic embryos of celery by control of gaseous compositions and other physical conditions

The celery (Tall Utah, Apium graveolens var. dulce) was adopted as a model system to investigate effective methods for enhancing embryogenesis. The focus was placed on attaining the optimum gaseous composition and the mixing intensity and viscosity of culture systems. The mixing intensity was investigated by using creased flasks and evaluated by shear force index (SFI). By using different closures with different ventilation coefficients, K w, to adjust the gaseous composition in the flasks, it was found that embryo frequency (EF) increased with high or low ethylene concentration in the headspace of the culture flasks. Cultivating the embryogenic cells under controlled ethylene concentrations depressed EF at ethylene concentrations higher than 0.08 ppm. Moreover, 0.1 μM of CoCl₂ remarkably repressed the ethylene formation in the culture medium and enhanced EF. and around 3% and 30%, respectively, favored EF. For non-creased flasks, addition of CMC (carboxymethyl cellulose) decreased the EF and DO level of the medium, due to lowered mass transfer rate attributed to high viscosity. For creased flasks, supplementation of 0.4 g/ 100 ml CMC reduced the ill-effects of high shear stress and maximized EF (7.22%). Our results are potentially applicable to other species and large-scale embryo production systems.     

Gill function in an elasmobranch

Summary Highly efficient oxygen uptake in elasmobranchs, as indicated by frequent excess of over has previously been ascribed to the operation of multicapillary rather than counter-current gas exchange by the gills. Analysis of models shows that, at maximum efficiency, a multicapillary system cannot account for values of greater than . In Port Jackson sharks Heterodontus portusjacksoni) commonly exceeds , which indicates the operation of a functional counter-current at the respiratory surface. The anatomical basis of this counter-current is provided by the demonstration that a continuous flow of water passes between the secondary lamellae into septal canals and thence via the parabranchial cavities to the exterior.Queen Elizabeth II Fellow.     

A new technique for continuous measurement of the heat of fermentation

Summary A special temperature control system has been developed and applied to continuous measuring of the heat evolved during a fermentation process. In this system, the fermentation broth was overcooled by a given constant cooling water flow. The excess heat removed from the fermentor was then made up by an immersion electrical heater. The action of the temperature controller was precisely monitored as it varied in response to the amount of heat produced by the microbial activities.The technique was used for determining the heat evolution byEscherichia coli grown on glucose. The ratio between quantities of total heat release and total oxygen consumption has been determined to be 0.556 MJ/mol O₂.The newly developed technique can be employed as an online sensor to monitor the microbial activities of either aerobic or anaerobic fermentation systems.Symbols C_c Heat capacity of cooling water (MJ/kg · °C) - C_p Heat capacity (MJ/kg · °C) - I Current of immersion heater (A) - K Constant in Equation (2) (h) - K Constant in Equation (13) (m³ · h · °C/MJ) - Q_c Flow rate of cooling water (m³/h) - Heat of agitation (MJ/m³ · h) - Heat dissipated by the bubbling gas (MJ/m³ · h) - Heat removal by the action of controller (MJ/m³ · h) - Heat of fermentation (MJ/m³ · h) - Heat loss to the surroundings (MJ/m³ · h) - Q_pass Constant average power dissipated by the immersion heater (MJ/m³ · h) - Fluctuating power dissipated by the immersion heater (MJ/m³ · h) - Power dissipated by the immersion heater (MJ/m³ · h) - T Temperature of fermentation broth (°C) - Constant average temperature of fermentation broth (°C) - Fluctuating temperature of fermentation broth (°C) - T_a Temperature of the ambient air (°C) - T_c Inlet temperature of cooling water (°C) - U₁A₁ Specific heat transfer coefficient for determination of heat loss to the surroundings (MJ/m³ · h · °C) - U₂A₂ Specific heat transfer coefficient for cooling surfaces (MJ/m³ · h · °C) - U₃A₃ Constant in Equation (16) (MJ/m³ · h · °C) - V Voltage of immersion heater (V) - V_L Liquid volume (m³) - OUR Oxygen uptake rate (mol O₂/m³ · h) Greek Letters H_fo The ratio between the total heat release and the total oxygen uptake (MJ/mol O₂) - _c Density of cooling water (kg/m³) - Time constant defined in Equation (6) (h) - _iM_iC_pi Heat capacity of system components (fermentation broth + fermentor jar + stainless steel) (MJ/m³ · °C)