Lifetime Reproductive Output Research Articles

The influence of diet on foraging habitat models: a case study using nursing Antarctic fur seals

Predictable sources of food underpin lifetime reproductive output in long lived animals. The most important foraging areas of top marine predators are therefore likely to be related to environmental features that enhance productivity in predictable spatial and temporal patterns. Even so, although productive areas within the marine environment are distributed patchily in space and time, most studies assess the relationships between feeding activity and proximate, not long term, environmental characteristics. In addition, individuals within a population may exploit different prey types, and these are often associated with different hydrographic features. Until now, models attempting to associate core foraging areas (CFAs) of marine predators with the environmental characteristics of those areas have not considered the diet of individual animals, despite the influence this could have on these relationships. We used bathymetry and multi‐year (n=24) mean sea surface temperature and variability as predictors of CFAs of lactating Antarctic fur seals Arctocephalus gazella at Heard Island. The effect of prey types on the predictability of these models was explored by matching diet and foraging trip data of individual seals (n=40 seals, n=1 trip each). Differences in diet between seals were mirrored by their spatial behaviour. Foraging strategies differed both between and within groups of seals consuming different diets. Long‐term environmental parameters were useful for predicting the foraging activity of seals that consumed a single prey type with relatively specific habitat preferences, but not for those that consumed single or multiple prey types associated with more varied habitats. Ignoring individual variation in predator diet probably contributes to the poor performance of foraging habitat models. These findings highlight the importance of incorporating individual specialization in foraging behaviour into ecological models and management of predator populations.

Shortened lifespan: another cost of fish-predator avoidance in cladocerans?

Life-history theory predicts adaptive shifts in response to size-selective predation, namely earlier reproduction, smaller age/size at maturity, and higher relative investment into reproduction. Such shifts should bring about reduced lifespan of potential prey. We tested this prediction in life-table experiments with clones of Daphnia hyalina and Diaphanosoma brachyurum, two species of contrasting anti-predatory strategies. The clones were derived from seven lakes of different trophy and held in water with and without fish kairomone, under standard laboratory conditions. Exposure to the kairomone caused a decrease in age of first reproduction and an increase in early-life reproductive effort but also an about 20% decrease of longevity in both species. Although shortened lifespan did not result in significant decrease in fitness of the tested species (in terms of lifetime reproductive output) it should be taken into account in considerations of costs and benefits of inducible defenses in cladocerans.

Predicting Life-History Traits for Female New Zealand Sea Lions, Phocarctos hookeri: Integrating Short-Term Mark-Recapture Data and Population Modeling

The trade-off between survival and reproduction by individuals is central to understanding life-history parameters of a species. Few mammal species have life-history information from long-term research. Instead, demographic models are commonly utilized to investigate an individual’s life-history strategy, species dynamics, and population trends. This research investigates age-related survival and reproductive performance of adult female New Zealand (NZ) sea lions (Phocarctos hookeri), using multi-state mark-recapture data from known-age branded individuals over five years. The mark-recapture analysis was integrated with a population model to predict the lifetime reproductive output of female NZ sea lions. The integration of an analysis of short-term datasets with population modeling allows for the prediction of life-history parameters of long lived animals when long-term information is not available. While such approaches involve some caveats, it provides a framework for investigating population dynamics and is preferential to unsubstantiated assumptions. This technique can lead to better design and implementation of conservation management for long lived species.

Lucilia sericata strain from Colombia: Experimental Colonization, Life Tables and Evaluation of Two Artifcial Diets of the Blowfy Lucilia sericata (Meigen) (Diptera: Calliphoridae), Bogotá, Colombia Strain

The objective of this work was to establish, under experimental laboratory conditions, a colony of Lucilia sericata, Bogotá-Colombia strain, to build life tables and evaluate two artificial diets. This blowfly is frequently used in postmortem interval studies and in injury treatment. The parental adult insects collected in Bogotá were maintained in cages at 22°C±1 average temperature, 60%±5 relative humidity and 12 h photoperiodicity. The blowflies were fed on two artificial diets that were evaluated over seven continuous generations. Reproductive and population parameters were assessed. The life cycle of the species was expressed in the number of days of the different stages: egg = 0.8±0.1, larvae I = 1.1±0.02, larvae II = 1.94±0.16, larvae III = 3.5±0.54, pupae = 6.55±0.47, male adult = 28.7±0.83 and female adult = 33.5±1.0. Total survival from egg stage to adult stage was 91.2% for diet 1, while for diet 2 this parameter was 40.5%. The lifetime reproductive output was 184.51±11.2 eggs per female. The population parameters, as well as the reproductive output of the blowflies that were assessed, showed relatively high values, giving evidence of the continuous increase of the strain over the different generations and making possible its maintenance as a stable colony that has lasted for more than two years.

The evolution of senescence in multi-component systems

Actuarial senescence is characterized by an increase in mortality rate with increasing chronological age. The reliability theory of senescence proposes that organisms' vital functions can be modelled as a suite of damageable, irreplaceable elements (typically genes or their products) that protect their bearer from condition-dependent death so long as at least one of the elements remains intact. Current incarnations of the reliability theory of senescence are continuous-time models with no explicit evolutionary component. Here, we use elementary probability theory and evolutionary dynamics analysis to derive a discrete-time version of the reliability theory of senescence. We include three variations on this theme: the 'Series' model in which damage to any of n elements results in death, the 'Parallel' model, in which damage accumulates in random order and damage to all n elements results in death, and the 'Cascade' (multi-stage) model, which is like the Parallel model, except the irreparable damage necessarily follows a strict sequence. For simplicity, we refer to the state of having multiple elements as 'redundancy', but this does not imply that the elements are necessarily identical. We show that redundancy leads to actuarial senescence in the Parallel and Cascade models but not in the Series model. We further demonstrate that in the Parallel and Cascade models, lifetime reproductive output (a potential proxy for fitness in populations with discrete generations) is a positive but decelerating function of redundancy. The positive nature of the fitness function leads to the prediction that redundancy and senescence should evolve from non-redundant, non-senescing ancestral populations; however, the deceleration of the fitness function leads to the prediction that this evolution towards increased redundancy will eventually be limited by mutation-selection balance. Using evolutionary dynamics analysis involving the discrete-generation quasispecies equation, we confirm these two predictions. Finally, we show that a population's equilibrium redundancy is sensitive to the environmental conditions that prevailed during its evolution, such as the rate of extrinsic mortality.

Differential reproductive strategies of two bivalves in the Dutch Wadden Sea

Cerastoderma edule and Mya arenaria are two common bivalve species in European waters. Longevity and maximum size are much greater in the latter species. Because comparison of species life-history strategies states that a long life span (i.e. high annual survival) generally goes with lower fecundity, we hypothesise that reproductive output would be lower in M. arenaria than in C. edule. In the present paper, we studied the reproductive strategies of these two species in an intertidal and a subtidal area of the western Dutch Wadden Sea, by following seasonal changes in absolute and relative weights of somatic and gonadal tissues in these bivalves. Starting of spawning was similar in the two species, around May, except for intertidal M. arenaria, which initiated spawning in August. Individual energy investment in reproduction was similar for the two species but, unlike M. arenaria, C. edule spawned completely, releasing all energy of gonadal mass in the form of gametes. Mya arenaria used the gonad not only for reproduction but also for storage. In the intertidal area, we found a trade-off between longevity and reproduction, i.e. maximum reproductive output (expressed as a proportion of body mass) was higher in C. edule than in M. arenaria. However, since body size is larger and life span longer in M. arenaria than in C. edule, mean lifetime reproductive output per individual must be higher in the first than in the latter. Based on the differences in reproductive strategies of these two species, we hypothesise that the negative effects of warming climate on bivalve population dynamics in the Wadden Sea will be stronger in C. edule than in M. arenaria.

Growth, mortality, and longevity of the Egyptian tortoise Testudo kleinmanni Lortet, 1883

Both demography and population regulation play an important role in the theory of sustainable exploitation and conservation of threatened taxa, such as terrestrial Chelonia. Here, we show and discuss some dynamic aspects of Testudo kleinmanni using modal progression analysis of length compositions. Although the Testudinata physiology is very different from that of fish, their growth model conforms to the Von Bertalanffy growth model. We observed a maximum of three age classes for both juveniles and females, and four classes for males. No appreciable between-sex differences were found in growth patterns, except for the diverse asymptotic length. Females should be subject to a strong sexual selection to quickly reach a large size in order to optimize lifetime reproductive output. The T. kleinmanni male size could be driven by predation escape and by easy accessibility to females, rather than by fighting for them. Thus, male reproductive success increases with the ability to fertilize females and female reproductive success increases with the ability to produce eggs, creating a large divergence in the context of selection between sexes. Different selective (synergetic or antagonistic) forces would appear to favor divergence in size between sexes. Additional properties found in the study regard the elevated mortality rate, probably also due to the human impact (poaching), and the relatively high longevity (26 and 22 years for females and males, respectively). Dynamics studies are useful for planning in situ activities of monitoring the population status, and could have a role in introducting programs and in control of reintroduced individuals during a restocking project.

Lifetime reproductive output over two generations in patients with psychosis and their unaffected siblings: the Uppsala 1915–1929 Birth Cohort Multigenerational Study

Schizophrenic patients have fewer offspring than the general population but it is unclear whether (i) this persists for more than one generation, (ii) the reduced fertility is compensated by increased fertility in unaffected relatives, (iii) sociodemographic factors confound or interact with the association, and (iv) patients with affective psychosis have a similar fertility disadvantage. This study measured biological fitness over two generations in patients with schizophrenia or affective psychosis, and their unaffected siblings. We conducted a historical cohort study using a Swedish birth cohort of 12 168 individuals born 1915-1929 and followed up until 2002. We compared biological fitness over two generations in patients with schizophrenia (n=58) or affective psychosis (n=153), and their unaffected siblings, with the population, adjusting for a range of sociodemographic variables from throughout the lifespan. Patients with schizophrenia had fewer children [fertility ratio (FR) 0.42, 95% confidence interval (CI) 0.29-0.61] and grandchildren (FR 0.51, 95% CI 0.33-0.80) than the population. Some of this reduction was related to lower marriage rates in schizophrenic patients. The unaffected siblings of schizophrenic patients showed no evidence of any compensatory increase in fitness, but there was a trend towards enhanced fertility among the offspring of schizophrenia patients. Patients with affective psychosis and their relatives did not differ from the general population on any fertility measure. Schizophrenia, but not affective psychosis, is associated with reduced biological fertility; this disadvantage is partly explained by marital status and persists into the second generation.

Going against the flow: effects of non-Gaussian dispersal kernels and reproduction over multiple generations

Our previous studies (Byers & Pringle 2006; Mar Ecol Prog Ser 313:27-41, Pringle & Wares 2007; Mar Ecol Prog Ser 335:69-84) discuss when populations and gradients in allele fre- quency can persist in the presence of a mean downstream dispersal of propagules from the parents' location. These studies assume that reproduction is uniform over the lifetime of the adult and that the larval dispersal kernel is 'nearly' Gaussian in shape. These results are extended in the present study to include variable reproduction over the lifetime of an adult and non-Gaussian dispersal kernels. It is found that persistence is governed by the lifetime reproductive output of the adults. The impact of non-Gaussian dispersal kernels is quantified in terms of the excess kurtosis of the dispersal kernel.

Life span, reproductive output, and reproductive opportunity in captive Goeldi's monkeys (Callimico goeldii)

In the absence of long-term field studies, demographic and reproductive records from animals housed in zoos and research laboratories are a valuable tool for the study of life history variables relating to reproduction. In this study, we analyzed studbook records of more than 2,000 individuals born over a 40-year period (1965-2004) to describe life history patterns of captive Goeldi's monkeys (Callimico goeldii) housed in North America and Europe. Using Kaplan-Meier survival analysis methods, we found the mean life span to be 5.5 years. The rate of infant mortality, defined as death before 30 days, was approximately 30%, with European animals being more likely to survive infancy than North American animals. When individuals surviving at least 1.5 years are considered, lifetime reproductive output averaged 3.5 offspring, yet more than one-third of individuals did not produce any offspring. Using a smaller dataset of individuals with known pairing histories, we developed a measure of opportunity for reproduction (OFR), which represented the total time an individual was known to be housed with a potential mate. For both sexes, we found that the correlation between OFR and number of offspring produced was much higher than the correlation between life span and number of offspring produced. This result highlights the importance of taking into account an individual's OFR. As a whole, our findings help characterize the life histories of captive Goeldi's monkeys and emphasize the impact management practices may have on reproductive success.

Overexpression of Glucose-6-phosphate Dehydrogenase Extends the Life Span of Drosophila melanogaster

The redox state of tissues tends to become progressively more prooxidizing during the aging process. The hypothesis tested in this study was that enhancement of reductive capacity by overexpression of glucose-6-phosphate dehydrogenase (G6PD), a key enzyme for NADPH biosynthesis, could protect against oxidative stress and extend the life span of transgenic Drosophila melanogaster. Overexpression of G6PD was achieved by combining a UAS-G6PD responder transgene at one of four independent loci with either a broad expression (armadillo-GAL4, Tubulin-GAL4, C23-GAL4, and da-GAL4) or a neuronal driver (D42-GAL4 and Appl-GAL4). The mean life spans of G6PD overexpressor flies were extended, in comparison with driver and responder controls, as follows: armadillo-GAL4 (up to 38%), Tubulin-GAL4 (up to 29%), C23-GAL4 (up to 27%), da-GAL4 (up to 24%), D42-GAL4 (up to 18%), and Appl-GAL4 (up to 16%). The G6PD enzymatic activity was increased, as were the levels of NADPH, NADH, and the GSH/GSSG ratio. Resistance to experimental oxidative stress was enhanced. Furthermore, metabolic rates and fertility were essentially the same in G6PD overexpressors and control flies. Collectively, the results demonstrate that enhancement of the NADPH biosynthetic capability can extend the life span of a relatively long-lived strain of flies, which supports the oxidative stress hypothesis of aging.

Optimal development time in pelagic copepods

According to life history theory, development times have evolved to an optimum solu- tion of the trade-offs between rapid development and high fecundity: rapidly developing individuals have a higher chance of surviving to maturity than do slowly developing ones, but they mature at a smaller size and, thus, have lower fecundity. Here we develop a model to predict egg-to-adult times in marine pelagic marine copepods that maximizes fitness, quantified either as lifetime reproductive output or population growth rate. The model is tested against global data sets of development time as a function of temperature. The optimum development time increases when the enhancement of fecundity with delayed maturation increases, and it decreases with increasing juvenile mortality. Our predictions match the patterns in nature for broadcast spawners, for which most data are available, but appear to underpredict development times for sac-spawning copepods. The partial uncoupling of growth and development rates in copepods demonstrated in laboratory experiments, the substantial variation in age at maturity observed in copepod field populations, as well as the fair match of pre- dicted and observed development times together suggest that development rates in marine pelagic copepods are tuned to the prevailing predation mortality. Observations in freshwater zooplankton suggest that life histories may be adapted to local conditions of growth and predation mortality. The striking lack of similar observations in marine zooplankton calls for work to explore the significance of environmental modulation of life histories of marine zooplankton. Possible future research avenues are proposed, including studies of the effects of seasonal environments, variable presence of preda- tors and the effects of sexual selection on life histories of marine planktonic copepods.

Variation in offspring sex ratio among individual Weddell seal (Leptonychotes weddellii) females of different quality

The Trivers–Willard model predicts that in polygynous species, superior-quality females will maximize their fitness by producing male offspring. Using a sample of 1,780 Weddell seal (Leptonychotes weddellii) pups recorded over 31 years, we investigated relationships between offspring sex ratio and maternal age, reproductive experience, an index of maternal lifetime reproductive output, and annual environmental variations. We found evidence that females with higher index of lifetime reproductive output were more likely to produce male than female offspring but found only weak evidence that large-scale environmental variations influenced sex ratios. Our results suggest that mothers manipulate offspring sex to maximize their own fitness, and inherent maternal quality may influence offspring sex. These findings support the Trivers–Willard sex-allocation model.

Biphasic growth in fish II: Empirical assessment

In [Quince, et al., 2008. Biphasic growth in fish I: Theoretical foundations. J. Theor. Biol., doi: 10.1016/j.jtbi.2008.05.029], we developed a set of biphasic somatic growth models, where maturation is accompanied by a deceleration of growth due to allocation of energy to reproduction. Here, we use growth data from both hatchery-raised and wild populations of a large freshwater fish (lake trout, Salvelinus namaycush) to test these models. We show that a generic biphasic model provides a better fit to these data than the von Bertalanffy model. We show that the observed deceleration of somatic growth in females varies directly with gonad weight at spawning, with observed egg volumes roughly 50% of the egg volumes predicted under the unrealistic assumption of perfectly efficient energy transfer from somatic lipids to egg lipids. We develop a Bayesian procedure to jointly fit a biphasic model to observed growth and maturity data. We show that two variants of the generic biphasic model, both of which assume that annual allocation to reproduction is adjusted to maximise lifetime reproductive output, provide complementary fits to wild population data: maturation time and early adult growth are best described by a model with no constraints on annual reproductive investment, while the growth of older fish is best described by a model that is constrained so that the ratio of gonad size to somatic weight ( g) is fixed. This behaviour is consistent with the additional observation that g increases with size and age among younger, smaller breeding females but reaches a plateau among older, larger females. We then fit both of these optimal models to growth and maturation data from nineteen wild populations to generate population-specific estimates of ‘adapted mortality’ rate: the adult mortality consistent with observed growth and maturation schedules, given that both schedules are adapted to maximise lifetime reproductive output. We show that these estimates are strongly correlated with independent estimates of the adult mortality experienced by these populations.

Biphasic growth in fish I: Theoretical foundations

We develop the theory of biphasic somatic growth in fish using models based on the distinction between pre- and post-maturation growth and an explicit description of energy allocation within a growing season. We define a ‘generic biphasic’ (GB) model that assumes post-maturation growth has a von Bertalanffy (vB) form. For this model we derive an explicit expression for the gonad weight/somatic weight ratio ( g) which may either remain fixed or vary with size. Optimal biphasic models are then developed with reproductive strategies that maximise lifetime reproductive output. We consider two optimal growth models. In the first (fixed g optimal), gonad weight is constrained to be proportional to somatic weight. In the second (variable g optimal) model, allocation to reproduction is unconstrained and g increases with size. For the first of these two models, adult growth in a scaled measure of length has the exact vB form. When there are no constraints on allocation, growth is vB to a very good approximation. In both models, pre-maturation growth is linear. In a companion paper we use growth data from lake trout ( Salvelinus namaycush) to test the bioenergetics assumptions used to develop these models, and demonstrate that they have advantages over the vB model, both in quality of fit, and in the information contained in the fitted parameters.

Postfledging parental effort in barn swallows: evidence for a trade-off in the allocation of time between broods

In altricial birds, parental care after fledging should be terminated as soon as the benefits from deserting the young outweigh those from continuing parental care. In seasonal environments, multiple breeders may trade off fitness benefits of the time invested in the care of fledglings of one brood against the benefits of an advanced start of the subsequent clutch. Using radiotelemetry of 494 fledglings in 114 families (of 88 pairs) of the double-brooded barn swallow, Hirundo rustica, we found that, irrespective of the offspring quality, the duration of postfledging parental care was shorter for first broods than for single and second broods. A shorter duration of postfledging care was associated with a steeper decline in feeding rates after fledging. Postfledging parental care also depended on the pair's breeding season length and on the phenotypic quality of the female parent. We suggest that pairs that start other reproductive bouts in the same season and pairs with high chances of survival to subsequent years reduce parental effort in the current brood to maximize lifetime reproductive output. The results support the hypothesis that multibrooded birds adjust the duration of postfledging care in relation to future reproductive attempts because of an intraseasonal trade-off in the allocation of time between successive broods.

Geographic variation in adult survival and reproductive tactics of the mosquito Aedes albopictus.

Climate differences across latitude can result in seasonal constraints and selection on life history characters. Since Aedes albopictus (Skuse) invaded North America in the mid-1980s, it has spread across a range of approximately 14 degrees latitude and populations in the north experience complete adult mortality due to cold winter temperatures that are absent in the south. Life table experiments were conducted to test for differences in the adult survival and reproductive schedules of Ae. albopictus females from two populations from the northern (Bloomington, IN [BL] and Manassas, VA [VA]; approximately 39 degrees N) and southern (Tampa, FL and Fort Myers, FL; approximately 27-28 degrees N) extremes of the species distribution in North America Regardless of population origin, age-specific hazard rate increased with reproductive output and decreased with number of bloodmeals. Larger females took fewer bloodmeals, and they had greater hazard rates than did smaller females. There were no consistent differences between northern versus southern populations in resource allocation between reproduction and maintenance, reproduction over time, and reproductive investment among offspring, suggesting that latitudinal variation in climate is probably not a main selective factor impinging on adult mortality and reproductive schedules. One possible effect of climate on geographic differences in life history was detected. BL had lower survivorship, lower lifetime reproductive output, and lower adult reproductive rate than did all other populations. This result may be an indirect result of lower egg survivorship due to the severity of winter in BL compared with other populations, including VA at approximately the same latitude. Such a scenario may make the BL population more prone to extinction, irregularly recolonized from more favorable sites, and thus more susceptible to founder effects, genetic drift, and inbreeding, resulting in lower mean values of fitness-related traits.

Nuptial gifts fail to resolve a sexual conflict in an insect.

BackgroundBecause of the potential benefits to individuals of saving investment for future mating opportunities, there is conflict between mates over most aspects of reproduction. Males of many species transfer compounds in the ejaculate that manipulate female reproductive physiology to increase male reproductive success. These seminal compounds are often associated with direct and/or indirect costs to females. In contrast, in some species ejaculates also contain nutrients used by females for somatic maintenance and increased reproductive output. In general, the extent to which male seminal components are detrimental or beneficial to females is poorly understood, and interactions between seminal compounds with different effects have been almost completely neglected. Here we examine the impact of male receptivity-suppressing factors and nutrient donations on female longevity and lifetime reproductive output in the bushcricket Requena verticalis.ResultsWe show that receiving multiple ejaculates reduces longevity in female R. verticalis, indicating a cost of male derived receptivity-suppressing compounds. Consumption of male nutrient donations does not appear to ameliorate this longevity cost, and there was no effect of nutrient provisioning on female lifetime fecundity.ConclusionThese results indicate that nutrient provisioning does not provide a resolution to sexual conflict over female receptivity in this bushcricket species.

Variation in age at first reproduction of male Japanese fluvial sculpin induced by the timing of parental reproduction

The relationships between age and size at reproduction and lifetime reproductive output of male Japanese fluvial sculpin Cottus pollux were estimated by a mark‐recapture study. Although all males were physiologically capable of breeding at age 2 years, age at first successful reproduction varied amongst individuals. Males with delayed reproduction had lower net reproductive rate than males that bred at age 2 years on average suggesting that age at first reproduction was a conditional strategy. Males that delayed reproduction were significantly smaller at age 1 and 2 years than males that bred at age 2 years. Despite no significant difference in body size of hatched yolk‐sac larvae between the early and late phase of the breeding season, by May of the first year of life, progeny from nests in the early phase had hatched earlier and were larger than those from the nests in the late phase. The results suggested an important effect of timing of reproduction of parents on the growth and subsequent age at first reproduction of their progeny.

Effects of feed composition on life history developments in feed intake, metabolism, growth and body composition of European eel, Anguilla anguilla

To examine the effect of feed composition on changes in feed intake and subsequent feed utilization with age, five populations of European eel, with an average initial body weight of 5 g each fed a different diet, were monitored for 302 d. The five feeds differed in their content of crude protein (33–63% DM), crude fat (6–28% DM) and calculated carbohydrates (NFE; 15–42% DM) such that five levels of digestible protein/digestible energy (DP/DE) were realised: 13, 16, 21, 28 and 29 g MJ − 1 . At three points in time, with three size groups, nitrogen and energy balance studies were conducted in which next to feed intake and growth also digestibilities of dry matter, protein, fat, NFE and energy as well as O 2 consumption and NH 4–N excretion were measured. Due to the distinct life history of the semelparous, in the present study predominantly male eel, a well-defined goal in terms of mature size and composition could be inferred, presumably to maximize their lifetime reproductive output. In order to reach this goal the animal needs to survive and to grow and voluntary feed intake of the eels could be adequately described with the feed intake model ‘eating to requirements subject to constraints’, where voluntary feed intake is considered to originate from a requirement for maintenance (survival) and a requirement for growth. Live weight gain is almost completely based on protein deposition (PD) and eels, like other animals, strive to reach a genetically determined growth potential (PDmax) thought to be driven by the difference from the mature protein mass (Ptmax). Body lipid content increases with size and varied with diet from a minimum of 25% at high DP/DE ratios to a maximum of 33% at low DP/DE ratios, at body weights of 130–140 g. Preferable allocation of dietary protein to PD (protein sparing action of non-protein energy) was confirmed as marginal efficiency of protein utilization increased with decreasing DP/DE ratio from 0.29 to 0.54. Marginal energetic efficiency of PD, k p was 0.54 and marginal energetic efficiency of LD, k f varied from 0.67, indicating de novo lipid synthesis (from dietary protein) at high DP/DE ratios, to 0.93, indicating direct lipid synthesis (from dietary lipid) at low DP/DE ratios. Marginal efficiencies did not differ from those of other fish or other farm animals. Differences between fish species in feed intake and utilization of feeds differing in macronutrient composition, as well as life history developments in feed intake and feed utilization are therefore based on differences in growth rate, in turn with mature weight (Ptmax), and body composition (LD/PD ratio).