A comparison between aerobic growth of Bacillus licheniformis in continuous culture and partial-recycling fermentor,with contributions to the discussion on maintenance energy demand

Energy costs of biomass synthesis are relatively higher at low than at high specific growth rates () because of an increased protein content of the cell and increased costs of protein synthesis as such at low values. A comparison of aerobic, glucose limited cultures of Bacillus licheniformis in a chemostat and in a partial-recycling fermentor indicated that pulse-wise nutrient addition increased the maintenance energy demand (m). In the chemostat experiments, we also found a striking deviation from linearity between substrate consumption and , with large implications for the maintenance coefficient. The deviation is mainly due to a large shift in metabolic carbon flows at specific growth rates between 50 and 100% of _max. At those growth rates, uncoupled growth occurs, presumably as a necessary condition for faster growth, since uncoupling results in a faster energysupply for biosynthetic purposes.The maintenance coefficient as determined by chemostat studies should be regarded as a compounds parameter, constituted of maintenance energy demands like ppGpp accumulation, variable costs of mRNA and protein accumulation, kinetic proofreading etc. and influenced by fermentor operation parameters like the substrate addition rate; moreover, both constancy of m and a linear relation between m and appear quite unlikely.     

Relationship between substrate inhibition and maintenance energy ofChlamydomonas reinhardtii in heterotrophic culture

The relationship between substrate inhibition and maintenance energy ofChlamydomonas reinhardtii grown heterotrophically on acetate was investigated. At low acetate concentrations (<0.4 g l^–1), where no inhibition of cell growth was observed, the cell growth yield and specific growth rate could be represented by the Pirt model, 1/Y=1/Y _g +m/ with a constant value of maintenance energy coefficient m. However, at high acetate concentrations (>0.4 g l^–1), inhibition of cell growth occurred, in which m became variable and dependent on the acetate concentration. A simple mathematical model was proposed to predict the actual maintenance energy coefficient m in the inhibited cultures and experimentally validated.Author for correspondence     

Very slow growth of Bacillus polymyxa: Stringent response and maintenance energy

Bacillus polymyxa grown in a recycling fermentor shows the same behavior previously observed with Escherichia coli: 3 successive growth phases. In the last 2 phases the growth rate is linear and the apparent maintenance energy demand rate and the molar growth yield are both independent of the specific growth rate, , and of the cells mass. The final phase of very slow growth is an indefinitely prolonged state of strong, stringent control, the regulatory system based on guanosine 3-diphosphate 5-diphosphate, and guanosine 3-diphosphate 5-triphosphate. The maximum cost of this stringent response is calculated to be 9% of the energy available to these energy-limited cells. There is a further energy cost contained in substantial amounts of DNA, RNA, and protein released from the cells during the latter 2 growth phases. The cost of production of these extra cellular anabolites ranges from 8–11% of the available energy.After a carbon-energy upshift in phase 3, the population growth rate immediately returned to that of early phase 2 growth, 50 h or more earlier.If maintenance energy is considered as energy expended by cells to maintain homeostasis, catabolic capacity, or anabolic potential, then the cost of stringent control — which preserves the fidelity of protein synthesis in slowly growing cells — must be considered a maintenance energy cost.Abbreviations GPR glucose provision rate - F_R medium flow rate - S_R substrate concentration - V_F fermentor volume - F_S filtrate removal rate - ppGpp guanosine 3-diphosphate 5-diphosphate - pppGpp guanosine 3-diphosphate 5-triphosphate     

Metabolic burden as reflected by maintenance coefficient of recombinant Escherichia coli overexpressing target gene

Metabolic burden as a consequence of overexpression of target gene in a recombinant strain of E. coli 1727 has been analyzed with respect to maintenance energy coefficient (m). The values of m for the host, uninduced recombinant and IPTG induced recombinant were determined to be 0.12, 0.17 and 0.32 g.g^-1.h^-1 respectively. Transient plasmid instability and nearly 33% fall in maximum specific growth rate were observed under conditions of enhanced requirements for maintenance energy.     

Interrelation between penicillin productivity and growth rate

Summary Fermentations were carried out in an 801 tower-loop reactor with pellets of Penicillium chrysogenum. The development of the inner structure of the pellets with regard to various fermentation conditions was observed by means of histological preparations of the pellets. Under conditions of energy-source-limitation mycelial tip growth and lysis of other mycelial parts exist simultaneously. Thus the net growth rate (formation rate of cell mass) is higher than the gross growth rate (multiplication rate of cell mass). Under conditions of nitrogen limitation, gross growth rate and net growth rate are identical. A very strict correlation between gross growth rate and penicillin production rate was found as long as sufficient oxygen supply could be maintained and carbon catabolite repression was avoided. The energy source requirement of the biomass can be described with the sum of three terms that correspond to gross growth, lysis compensation growth and maintenance.Symbols a Constant 1/l h - b Constant - K Decay rate constant for product 1/h - K ₁ Substrate inhibition constant g/l - K _op Controls saturation constant for oxygen g/l - K _p Saturation constant for substrate g/l - m Maintenance coefficient 1/h - ms Apparent maintenance coefficient 1/h - O Dissolved oxygen concentration g/l - P Product concentration g/l - p Exponent of O - q Specific productivity 1/h - S Substrate concentration g/l - t Time h - t ₁ Beginning of production phase h - t ₂ Time of pellet dissolution h - V Liquid volume of fermentation broth l - X Dry cell mass concentration g/l - Y Yield of dry cell mass from energy substrate - g Specific gross growth rate of biomass 1/h - l Specific lysis rate of cell mass 1/h - n Specific net growth rate of cell mass 1/h - p Maximum specific rate of product formation 1/h     

Maintenance energy demand affects biomass synthesis but not cellulase production by a mesophilicClostridium

Summary Secretion of cellulolytic activity by the mesophilClostridium strain C7 was studied while the bacterium underwent progressive carbon/energy starvation and the ensuing continuous decline in growth rate. In the slowest range of growth rates studied the organism was in full response to the global regulation imposed by guanosine 5, 3-bispyrophosphate (ppGpp). The exoenzymes of the cellulase complex were produced at the same volumetric rate whether or not the response was active. However, the volumetric rate of biomass synthesis was reduced 45% or more by the response. Energy necessary to maintain the ppGpp-regulated state (i.e., maintenance energy) was, therefore, diverted from energy going to synthesis of biomass but not from that going to exoenzyme synthesis, making the yield of cellulase activity per mole of carbon-energy substrate independent of growth rate and the exoenzyme complex produced from the substrate with equal efficiency at all growth rates. The primary consideration in improving exoenzyme productivity by bacteria with this type of energy distribution between secretion, growth, and maintenance is simply increasing yield per mole of carbon-energy substrate, with growth rate effects on yield a secondary and minimum concern.     

Eubacteria have 3 growth modes keyed to nutrient flow

Aerobic growth of Escherichia coli and Paracoccus denitrificans has been studied in chemostat, fed batch, and recycling fermentor modes under carbon and energy limitation. Two abrupt drops or discontinuities in molar growth yield, Y, have been found that occur over relatively short ranges in the value of specific growth rate.Before the first discontinuity, Y is constant and maximal. After the first discontinuity, at a doubling time of 33 h, Y becomes constant again and independent of until the second discontinuity appears at a doubling time of about 50 h, corresponding to a of about 0.014. At this point, Y drops to a lower value that is constant at doubling times longer than 100 h, corresponding to a of about 0.007.The second discontinuity is associated in Paracoccus with elevated levels of guanosine tetraphosphate (ppGpp) that impose stringent regulation as has been found previously with Bacillus and Escherichia species. It is thus likely that the stringent response generally occurs in bacteria in vivo at a doubling time of about 50 h. The cause of the first discontinuity is unknown. All experiments indicate that Pirt-type calculations relating , Y, and maintenance energy demand are no longer valid. In chemostat experiments, the intercept of the relationship between specific substrate utilization and specific growth rate is defined as maintenance. However, this intercept most probably is caused by stringent regulation at low dilution rates. Three regions of bacterial growth rates are defined by this study, corresponding to doubling times of 0.5 to 15 h, 33 to 50 h, and >100 h. Some growth behavior in each region is unique to that region.Abbreviations ppGpp guanosine 5 diP 3 diP - pppGpp guanosine 5 triP 3 diP - SPR substrate provision rate (mol/l h)     

The energetics of starvation and growth after refeeding in juveniles of three cyprinid species

Synopsis Experiments were conducted to monitor changes in body mass and metabolic energy expenditure before, during, and after periods of starvation in juveniles of three species of cyprinids: Leuciscus cephalus, Chalcalburnus chalcoides mento, and Scardinius erythrophthalmus. During the starvation period all fish lost weight at about the same rate and the total amount of oxygen consumed during an experimental period of 20 h was about 40% lower in the starved than in the fed groups. Upon refeeding, both mass specific maintenance; and routine rates of metabolism as well as relative growth rates increased rapidly, the peaks of these increases being directly proportional to the length of the starvation period. Maximum compensatory growth was observed after four weeks of starvation in C. chalcoides and S. erythrophthalmus, with relative growth rates reaching 30% d^-1 during the first measuring interval after refeeding. The pattern of time-dependent compensatory growth displayed by these fish is similar to the responses of a colonial hydroid in which the rate of catch-up growth increased with the amount of stress to which the animals had been exposed. The exact cost of compensatory growth cannot be calculated because oxygen consumption and growth were not measured simultaneously. However, on the basis of data and calculations reported by Wieser & Medgyesy (1990) it appears that compensatory growth, if fuelled by the metabolic power indicated by our measurements of oxygen consumption, would have to be about twice as efficient as normal growth in the related species Rutilus rutilus.     

The physiological ecology of two populations of Mytilus edulis L.

Summary Seasonal cycles in the rates of oxygen consumption, feeding, absorption efficiency and ammonia-nitrogen excretion in two populations of Mytilus edulis were measured in the field under ambient conditions and related to body size, the gametogenic cycle, the concentration of suspended particulate matter in the water and temperature. Relationships between the various physiological variables are also considered and protein and energy budgets estimated. Both the scope for growth and the relative maintenance cost were seasonally variable, demonstrating a minimum capacity for growth in the winter and a maximum capacity in the summer. In one population subjected to abnormally high temperatures in the winter the scope for growth was negative for four or five months between January and May. These population differences are discussed and the potential for using physiological integrations in intra-specific comparisons of fitness is identified.     

Temperature and food quality influences feeding behavior,assimilation efficiency and growth rate of arctic woolly-bear caterpillars

Summary The energy budget for feeding activity and growth of larval Gynaephora groenlandica was investigated on the tundra and in the laboratory. Larvae fed only in June when the buds and young leaves of Salix arctica, its principal host plant, contained the highest concentrations of macro-nutrients and total nonstructural carbohydrates (TNC). The mid-summer hiatus in larval feeding was coincident with an abrupt decline in the TNC content of leaves and a buildup of plant secondary metabolites in the leaves of S. arctica. Following cessation of feeding, the larvae remained concealed from the sun within crevices and vegetation mats. Growth rates of larvae incubated at 15 and 30°C were similar (4.7–5.0 mg/larva/day), but the assimilation efficiency at 15°C was four times greater (40%) than at 30°C. Growth rates were lowest at 5°C (0.22mg/larva/day) as was the assimilation efficiency (6.6%), because of the extended residence time of food in the gut. The high rate of ingestion and excretion at 30°C was caused by elevated maintenance metabolism. Changes in metabolic state influenced oxygen consumption, which was highest for feeding larvae (0.29 ml/g/h) and significantly lower for each, digesting, moving, starved larvae, and lowest for inactive larvae (0.06 ml/g/h). An influence of temperature and leaf quality on digestion rate and maintenance metabolism is the most likely cause of the feeding behavior pattern in G. groenlandica. The larvae may undergo voluntary hypothermia in order to avoid an energy, deficit resulting from high maintenance metabolism during mid-season when the energy content and food quality declines. The restriction of growth and development to a very short period prior to mid-summer may have contributed, to the extended 14-year life cycle of this species.     

The role of energy-spilling reactions in the growth ofKlebsiella aerogenes NCTC 418 in aerobic chemostat culture

When cell-saturating amounts of glucose and phosphate were added to steady state cultures ofKlebsiella aerogenes that were, respectively, glucose-and phosphate-limited, the organisms responded immediately with an increased oxygen consumption rate. This suggested that in neither case was glucose transport the rate-limiting process, and also that organisms must posses effective mechanisms for spilling the excess energy initially generated when a growth-limitation is temporarily relieved.Steady state cultures of mannitol- or glucose-limited organisms also seemingly generated energy at a greater rate than was required for cell synthesis since gluconate-limited cultures consumed oxygen at a lower rate, at each corresponding growth rate, than did mannitol- or glucose-limited cultures, and there-fore expressed a higherY _o value. Thus, mannitol- and glucose-limitations must be essentially carbon (and not energy) limitations. The excess energy generated by glucose metabolism is one component of maintenance and could be used at lower growth rates to maintain an increased solute gradient across the cell membrane, imposed by the addition of 2%, w/v, NaCl to the growth environment.The maintenance rates of oxygen consumption ofK. aerogenes also could be caused to increase by adding glucose discontinuously (drop-wise) to a glucose-limited chemostat culture, or by exchanging nitrate for ammonia as the sole utilizable nitrogen source.The significance of these findings to an assessment of the physiological factors circumscribing energy-spilling reactions in aerobic cultures ofK. aerogenes is discussed.     

Energy requirement for maintenance of the transmembrane potassium gradient in Klebsiella aerogenes NCTC 418: A continuous culture study

Molar growth yield studies on chemostat cultures of Thiobacillus neapolitanus grown in thiosulfate-minerals medium have confirmed earlier observations that the dry weight increased linearly with the dilution rate. The observed increase can be explained neither by a change in cell composition nor by the observed excretion of organic compounds. The increase of the molar growth yield over the full range of growth rates, that is also observed in other obligate chemolithotrophs, was not found in the facultatively chemolithotrophic Thiobacillus A2, grown on thiosulfate or formate. The interpretation of the results in terms of maintenance energy requirement is discussed. It is concluded that these results do not allow a mathematical treatment according to the empirical formula of Pirt.Abbreviation APS Adenosine-5-phosphosulfate     

Lactic acid production by batch fermentation of whey permeate: A mathematical model

Summary The batch fermentation of whey permeate to lactic acid was improved by supplementing the broth with enzyme-hydrolyzed whey protein. A mathematical model based on laboratory results predicts to a 99% confidence limit the kinetics of this fermentation. Cell growth, acid production and protein and sugar use rates are defined in quantifiable terms related to the state of cell metabolism. The model shows that the constants of the Leudeking-Piret model are not true constants, but must vary with the medium composition, and especially the peptide average molecular weight. The kinetic mechanism on which the model is based also is presented.Nomenclature K _i lactic acid inhibition constant (g/l) - K _pr protein saturation constant during cell growth (g/l) - K _pr protein saturation constant during maintenance (g/l) - K _s lactose saturation constant (g/l) - [LA] lactic acid concentration (g/l) - [PR] protein concentration (g/l) - [S] lactose concentration (g/l) - t time (h) - [X] cell mass concentration (g/l) - , fermentation constants of Leudeking and Piret - specific growth rate (l/h) - Y _{g, LA/S} acid yield during cell growth (g acid/g sugar) - Y _{m, LA/S} acid yield during maintenance (g acid/g sugar) - Y _x/pr yield (g cells/g protein) - specific sugar use rate during cell growth (g sugar/h·g cell) - specific sugar use rate during maintenance (g sugar/h·cell)     

Influence de la température sur la cinétique,de croissance et le coefficient de maintenance de Candida lipolytica cultivé sur n-alcane

The effect of growth temperature on the evolution of kinetic parameters and yields was determined for Candida lipolytica cultures with ntetradecane as substrate, in a temperature range of 18°C to 30°C, which is below the critical growth temperature in order to work only in the activation zone of these parameters.In such a culture limited by substrate transfer, growth rate depends on biological rates, related to microorganisms characteristics, and diffusional rates, related to mass transfer. The effect of temperature thus depends on the limiting step. The activation energy, calculated from exponential growth rate determinations is .When the activation energy is calculated from the maximal rate of cell production (determined at the growth curve's inflexion point), it's found to be E _X=71,200 J/mole in the 18°C–24°C range, and E _X=28,000 J/mole in the 24°C–30°C range. The latter one is characteristic of a diffusion-limited process. Above 24°C, growth is controlled by substrate-transfer, as physiological potentialities are preferentially increased with temperature than diffusional ones: 24°C is thus the transition temperature T _t from physiological to diffusional limitation.The apparent yield is almost constant, over the 18°C to 30°C temperature range, although maintenance coefficients are very dependent on temperature. The activation energies related to maintenance coefficients for alkane and oxygen respectively are and .The m _s/m_{O 2} ratio is about 3 (g/g), whereas that, for a strict oxidation reaction of n-tetradecane ought to be 3.47 (g/g). A satisfactory correlation, relating maintenance coefficients to the maximal growth rate of yeast, is given.

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Liste des symboles A constante de saturation de modèle de croissance(1) - B vitesse spécifique considérée - C substrat carboné ou oxygène (g/l) - E energie d'activation (J/mole) - S _m quantité de substrat consommée par maintenance au cours d'une fermentation discontinue (g) - O₂ quantité d'oxygène transférée au milieu de culture (g/l) - R rendement global de la fermentation - R rendement global de la fermentation - constante des gaz parfaits (J/mole K) - S concentration en substrat carboné (g/l) - T température de croissance (°K) - X concentration en biomasse (g/l) - Y rendement limite - m coefficient de maintenance (h^-1) - t duree de fermentation (h) - tømpérature de croissance (^o Celsius) - taux de croissance (h^-1) Indices 1 relatif à la température 1. - 2 relatif à la température 2 - c relatif au substrat carboné ou à l'oxygène - f relatif au temps final - i relatif au point d'inflexion - m maximum - mO₂ relatif au coefficient de maintenance sur l'oxygène - m _s relatif au coefficient de maintenance sur le substrat carboné - o relatif au temps initial - O₂ relatif à l'oxygène - s relatif au substrat carboné - t de transition - T relatif à la température de croissance T - U _m relatif au taux de croissance maximal - X relatif à la productivité maximale en biomasse     

Survival of Chromatium vinosum at low light intensities

The effect of low irradiation on the viability of Chromatium vinosum was investigated. Cultures were precultivated at 1,000 lux (=0.1/h). Then, before the substrate was depleted, illumination was changed to either complete darkness or about 30 lux. Previously, the latter light intensity had been found not to promote growth.The parameters assayed were viability, protein, bacteriochlorophyll, ATP, RNA, DNA, absorbance (E ₂₆₀) of the supernatant, and total anthron-positive material.The data show that irradiation insufficiently high to promote growth, results in viability percentages as high as 90% after 8 days, whereas cultures incubated in complete darkness are virtually dead by then. Neither in the light nor in the dark a degradation of protein or cell wall hexoses was observed. The RNA content also remained constant. However, particularly in the dark cultures DNA was found to decrease concomitant with increased E ₂₆₀ readings of the supernatant. It is considered unlikely that such essential macromolecules are degraded to serve the maintenance energy requirements. The ecological impact of the observations is discussed.Non-Standard Abbreviations PHB poly--hydroxybutyric acid - Bchl Bacteriochlorophyll     

Isolation and growth of a bacterium able to degrade nitrilotriacetic acid under denitrifying conditions

A Gram-negative bacterium was isolated from river sediment which was able to grow with nitrilotriacetic acid as a combined carbon, nitrogen and energy source in the absence of molecular oxygen using nitrate as the terminal electron acceptor. Batch growth parameters and mass balances are reported for growth under both aerobic and denitrifying conditions.The strain was characterized with respect to its substrate spectrum and other physiological properties. This denitrifying isolate is serologically unrelated to the comprehensively described Gram-negative obligately aerobic NTA-degrading bacteria all of which belong to the -subclass of Proteobacteria. Chemotaxonomic characterization, which revealed the presence of spermidine as the main polyamine and ubiquinone Q-8, excludes the new isolate from the phylogenetically redefined genus Pseudomonas and indicates a possible location within the -subclass of Proteobacteria close to, but separate from the genus Xanthomonas.     

The effect of temperature on the growth of A. niger in solid state fermentation

A kinetic model of solid state fermentation with temperature deactivation of microorganisms is presented. The experimental results of cultivation of Aspergillus niger on a mixture of wheat bran and beet pulp in temperature range from 26 °C to 40 °C were used to estimate the parameters of the model. The activation energies of growth, thermal deactivation and maintenance have been calculated.List of Symbols C _CX mol/g proportionality coefficient - E _d J/mol energy of activation for thermal deactivation - E _g J/mol energy of activation for growth - J _CO2 mol/gh carbon dioxide evolution rate - k _d h^–1 thermal deactivation constant - k _g h^–1 growth kinetic constant - k _x h^–1 net growth constant - m h^–1 maintenance coefficient - N _CO2 mol amount of carbon dioxide - N _{m, CO2} mol maximum amount of carbon dioxide generated by growth - t h time - X g dry biomass weight - X _m g maximum biomass weight - X dimensionless biomass weight - X _0,r g real mass of inoculum - X _0,a g apparent mass of inoculum - X ₀ dimensionless apparent mass of inoculum - dimensionless maintenance coefficient This work was supported by the Committee of Scientific Research under grant No 3 3401 91 02.     

Chemolithotrophic metabolism of the newly-isolated moderately thermophilic,obligately autotrophic Thiobacillus tepidarius

Thiobacillus tepidarius, isolated from the hot springs at Bath, Avon, UK, grew optimally at 43–45°C and pH 6.0–7.5 on thiosulphate or tetrathionate. In batch culture, thiosulphate was oxidized stoichiometrically to tetrathionate, with a rise in pH. The tetrathionate was then oxidized to sulphate, supporting growth and producing a fall in pH to a minimum of ph 4.8. The organism contained high levels of thiosulphate-oxidizing enzyme, rhodanese and ribulose bisphosphate carboxylase. It was obligately chemolithotrophic and autotrophic. In chemostat culture, T. tepidarius grew autotrophically with the following sole energy-substrates: sulphide, thiosulphate, trithionate, tetrathionate, hexathionate or heptathionate. Thiocyanate, dithionate and sulphite were not used as sole substrates, although sulphite enhanced growth yields in the presence of thiosulphate. Maximum specific growth rate on tetrathionate was 0.44 h^-1. True growth yields (Y _max) and maintenance coefficients (m) were calculated for sulphide, thiosulphate, trithionate and tetrathionate and observed yields at a single fixed dilution rate compared with those on hexathionate and heptathionate. Mean values for Y _max, determined from measurements of absorbance, dry wt, total organic carbon and cell protein, were similar for sulphide, thiosulphate and trithionate (10.9 g dry wt/mol substrate) as expected from their equivalent oxygen consumption for oxidation. Y _max for tetrathionate (20.5) and the relative Y _o values (as g dry wt/g atom oxygen consumed) for thiosulphate and all four polythionates indicated that substrate level phosphorylation did not contribute significantly to energy conservation. These Y _max values were 40–70% higher than any of those previously reported for obligately aerobic thiobacilli. Mean values for m were 6.7 mmol substrate oxidized/g dry wt·h for sulphide, thiosulphate and trithionate, and 2.6 for tetrathionate.Abbreviation PIPES Piperazine-N,N-bis(ethane sulphonic acid)     

Growth yields of Desulfotomaculum orientis with hydrogen in chemostat culture

Desulfotomaculum orientis (strain Singapore 1) was grown autotrophically with H₂+CO₂ and sulfate, thiosulfate or sulfite as electron acceptor in sulfide- and pH-controlled continuous culture. Under sulfate-limiting conditions real growth yields of up to 9.7 g cell dry mass per mol sulfate were obtained. Electron acceptor limitation resulted in the excretion of up to 14.5 mmol acetate per liter, formed by reduction of CO₂ with H₂. Acetate production was not coupled to an increase of growth yields: under hydrogen-limiting conditions only 1.6 mmol acetate per liter was produced, and even higher growth yields of up to 12,4 g cell dry mass per mol sulfate were obtained. With thiosulfate or sulfite as electron acceptor growth yields increased up to 17.9 g cell dry mass per mol electron acceptor. Growth yields were not simply correlated with the growth rate, and did not allow the determination of maintenance coefficients and the extrapolation to maximal yields at infinite growth rate (Y ^max). The maximal growth rates (_max) with sulfate and thiosulfate were 0.090 and 0.109 h^-1, respectively, if cells were grown continuously in sulfidostat culture under nonlimiting conditions.The net energy yield of sulfate reduction and the energy requirement for the activation of sulfate by Desulfotomaculum orientis are discussed.     

Influence of temperature and food availability on the ecological energetics of the giant scallop Placopecten magellanicus

Summary Gonad output, reproductive effort and residual reproductive value are greater in scallops (Placopecten magellanicus) from shallow water, where conditions are more favourable, than in scallops from deep water. Variation between years in these fitness correlates is also greater in shallow water scallops. High reproductive effort is associated with a greater reproductive cost, although in general reproduction in this species appears to be restrained, and only in older individuals from shallow water is there evidence of reckless reproductive behaviour. Reproductive effort increases as the animal grows older, but residual reproductive value is a maximum at age 4 years, after which it decreases with age owing to greater mortality. Localities vary in their suitability for scallop growth and reproduction, and at one Newfoundland site conditions in shallow water (10 m) are better than those in deep water (31 m). Scallops from the former have a greater fitness than those from the latter. The energy costs of reproduction have implications for life history variability in P. magellanicus, although for a considerable part of its lifetime the scallop is able to maximise gamete production in response to environmental factors (especially the food supply) without trading off reproduction against growth or maintenance.