Energy Investment In Reproduction Research Articles

Evolution Under Dietary Restriction Increases Reproduction at the Cost of Decreased Somatic Growth.

Dietary restriction (DR) is cited as the most reliable means of extending life span in a wide range of taxa, yet the evolutionary basis of the DR effect on life span remains unclear. The resource reallocation hypothesis proposes that the longevity-extending response of DR is adaptive and stems from the reallocation of resources from reproduction to somatic maintenance under food-limited conditions. However, if DR continues for multiple generations, such a response becomes maladaptive, and genotypes with higher fecundity should be selectively favored over genotypes with longer longevity. To test this prediction, we exposed replicate populations of the rotifer Brachionus dorcas, a model organism for aging and experimental evolution studies, to DR and ad-libitum (AL) diets for 100 days. During the selection experiment, AL-selected populations showed higher growth rates and mictic ratios than DR-selected populations. After approximately 27 asexual generations of selection, populations with a DR selection history had a higher net reproductive rate but lower body volume and ingestion rate in the absence of survival costs than populations with an AL selection history when they were assayed on an AL diet. Our results are inconsistent with the prediction that evolution on sustained DR increases reproduction and reduces life span, and show for the first time that sustained DR selects for clones with higher energy investment in reproduction but lower investment in somatic growth.

The effect of pregnancy on metabolic scaling and population energy demand in the viviparous fish Gambusia affinis.

Metabolism is a fundamental attribute of all organisms that influences how species affect and are affected by their natural environment. Differences between sexes in ectothermic species may substantially alter metabolic scaling patterns, particularly in viviparous or live-bearing species where females must support their basal metabolic costs and that of their embryos. Indeed, if pregnancy is associated with marked increases in metabolic demand and alters scaling patterns between sexes, this could in turn interact with natural sex ratio variation in nature to affect population-level energy demand. Here, we aimed to understand how sex and pregnancy influence metabolic scaling and how differences between sexes affect energy demand in Gambusia affinis (Western mosquitofish). Using the same method, we measured routine metabolic rate in the field on reproductively active fish and in the laboratory on virgin fish. Our data suggest that changes in energy expenditure related to pregnancy may lead to steeper scaling coefficients in females (b=0.750) compared to males (b=0.595). In contrast, virgin females and males had similar scaling coefficients, suggesting negligible sex differences in metabolic costs in reproductively inactive fish. Further, our data suggest that incorporating sex differences in allometric scaling may alter population-level energy demand by as much as 20-28%, with the most pronounced changes apparent in male-biased populations due to the lower scaling coefficient of males. Overall, our data suggest that differences in energy investment in reproduction between sexes driven by pregnancy may alter allometric scaling and population-level energy demand.

A Histopathological–Biochemical Health Assessment of Blue Mussel Mytilus edulis

Aquaculture of the blue mussel Mytilus edulis in the Gulf of Maine is a growing industry at a time when wild mussel populations are threatened by a rapidly changing ocean. Intertidal mussel beds have largely disappeared in the region raising concerns over the long-term viability of mussel farming. Histology and lipid fatty acid analysis were used to assess gametogenesis, energy investment, and pathology of farmed mussels collected twice monthly for three years in Casco Bay, ME. Energy investment in reproduction and storage differed significantly between years, suggesting interannual variability. Wet weight of fatty acids such as DHA and EPA corresponded to pre-spawning periods, when gonad tissue was most abundant. Overall, pathology assessment showed low levels of common pathogens, parasites, and cellular abnormalities. The survey did, however, reveal high levels of oocyte atresia, a probable indicator of physiological or environmental stress from unfavorable spawning conditions. In addition, the presence of the potentially damaging digenetic trematode Proctoeces maculatus was documented using histology, marking the northernmost detection in the Northwest Atlantic and a likely climate-driven range expansion. These trends may signal a challenging future for blue mussels in the Gulf of Maine. Forward-looking farm mitigation practices informed by these results should be developed to ensure future sustainability of this industry.

Body condition and energy content of shore crab Carcinus maenas in a temperate coastal system: temporal variability

The body condition of the shore crabCarcinus maenasin a temperate coastal system (western Dutch Wadden Sea) was followed over 14 mo. Fulton’s condition factorK, dry weight condition and the percentage of dry weight were determined as indirect indices, while bomb calorimetry was applied to obtain the energy density and total energy, as direct biochemical measures of condition. General linear models identified sex, size and season as relevant regressors explaining observed variance in crab condition, whereas colour morphotype effect was negligible. The seasonal pattern was consistent with the natural cycling in temperature and expected food availability, but the peak in body condition differed depending on the type of measure used: energy peaked in autumn, while morphometric condition was at its highest in winter, uncovering different latencies in the response of direct and indirect indices. Concordant with higher energy investment in reproduction, body condition and energy content of non-ovigerous females were higher than those of males, and egg-bearing females always had the lowest condition values. Energy content of adult females ranged from 16.37 ± 1.30 (winter 2013) to 19.83 ± 0.54 kJ g-1ash-free dry weight (AFDW) (autumn 2013), attaining 18.77 ± 1.22 kJ g-1AFDW prior to the onset of reproduction in 2012, while maximum energy density of ovigerous females (eggs excluded) was 16.49 ± 0.64 kJ g-1AFDW. Besides low correlation between indirect and direct indices, fluctuations were more pronounced in the energetic data. Therefore, the morphometric measures seem weak surrogates to estimate energy density of these crabs.

Reproduction and Seasonal Growth Estimates of the Black Goby, Gobius niger (Osteichthyes: Perciformes: Gobiidae) in the Southeastern Black Sea Region of Turkey

Seasonal growth pattern and reproductive biology of the black goby, Gobius niger, were studied in southern Black Sea between December 2012 and November 2013. The seasonal von Bertalanffy growth parameters, computed from ELEFAN, were estimated as L∞ = 11.90 cm TL, K = 0.701 year-1, C = 0.633, and WP = 0.492 for the Hoenig seasonal model and as L∞ = 12.14 cm TL, K = 0.680 year-1 for non-seasonal von Bertalanffy model. The size at sexual maturity (TL50) was estimated as 8.9 cm TL. The GSI ranged 0.60 in December and 8.57 in April (mean 3.82±0.818) for females and between 0.61 in October 2013 and 2.90 in May 2013 (mean 1.70±0.260) for males. The seasonal growth was pronounced probably due to reproduction activity which extended investment of energy in reproduction causing slower growth. The GSI development was related to photoperiod and the monthly variations of GSI values indicating that the intensive spawning occurred between April and August. During spawning time, the mean GSI value of females (6.04±1.177) was 2.9 times higher than the mean GSI values of males (2.08±0.423) (P < 0.05). The results of this study were offered as biological input parameters regarded as a reference for the conservation and management of the Black Sea stocks of the black goby.

Faecal parasites increase with age but not reproductive effort in wild female chimpanzees.

Energy investment in reproduction is predicted to trade off against other necessary physiological functions like immunity, but it is unclear to what extent this impacts fitness in long-lived species. Among mammals, female primates, and especially apes, exhibit extensive periods of investment in each offspring. During this time, energy diverted to gestation and lactation is hypothesized to incur short and long-term deficits in maternal immunity and lead to accelerated ageing. We examined the relationship between reproduction and immunity, as measured by faecal parasite counts, in wild female chimpanzees (Pan troglodytes schweinfurthii) of Kibale National Park, Uganda. While we observed higher parasite shedding (counts of eggs, cysts and larvae) in pregnant chimpanzees relative to cycling females, parasites rapidly decreased during early lactation, the most energetically taxing phase of the reproductive cycle. Additionally, while our results indicate that parasite shedding increases with age, females with higher fertility for their age had lower faecal parasite counts. Such findings support the hypothesis that the relatively conservative rate of female reproduction in chimpanzees may be protective against the negative effects of reproductive effort on health. This article is part of the theme issue 'Evolution of the primate ageing process'.

Do males and females respond differently to ocean acidification? An experimental study with the sea urchin Paracentrotus lividus

Seawater pH lowering, known as ocean acidification, is considered among the major threats to marine environment. In this study, post-spawning adults of the sea urchin Paracentrotus lividus were maintained at three pH values (8.0, 7.7, 7.4) for 60 days. Physiological, biochemical, cellular, behavioural and reproductive responses were evaluated in males and females. Significant differences between sexes were observed, with higher ammonia excretion and lower catalase activity in males. Respiration rate (after 21 days), catalase activity in gonads and total coelomocyte count showed the same increasing trend in males and females under low pH. Ammonia excretion, gonadosomatic index and lysozyme activity exhibited opposite responses to low pH, with an increasing trend in males and decreasing in females. Results demonstrated that exposure to low pH could result in different response strategies of male and female sea urchins at a physiological, biochemical and immunological level. Reduced female gonadosomatic index under low pH suggested decreased energy investment in reproduction.

Sex-specific structural and functional leaf traits and sun-shade acclimation in the dioecious tree Pistacia vera (Anacardiaceae).

In dioecious species, sex-related adaptive strategies, influenced by natural and sexual selection, allow each sex to meet the specific demands of reproduction. Differences in ecophysiological traits between males and females may rely on innate differences in secondary sex traits such as structural and functional leaf traits. We tested structural sexual leaf dimorphism in Pistacia vera L. and the intersexual differences in sun-shade acclimation processes expected from the different adaptive strategies of males and females. Fifteen structural and functional leaf traits were compared in 50-year-old trees between females with low fruit load and males under sun and shade conditions. Despite the low additional energy investment in reproduction in females, remarkable sex effects in leaf structure and function were observed. Male trees had smaller leaves with significantly lower total conducting petiole area (TCA) and higher stomatal density, water use efficiency and concentration of phenolic compounds; females had larger leaves with greater thickness, leaf mass per area, TCA and maximum photosynthetic capacity per area (Amax,a). The higher Amax,a and stomatal conductance of female leaves were associated with their ~20-fold higher TCA compared with male trees. Females seem to invest more in high xylem efficiency and rates of C gain; males invest more in defence-protection. Sun-shade plastic responses were sex- and trait-specific, but the plasticity assessment indicated that both sexes have evolved an almost equal degree of phenotypic plasticity that allows them to perform optimally under varying environmental conditions. However, the trait-specific differences indicate that each sex displays a different strategy of optimisation.

Geographical variation and local environment effects in the reproductive output and fecundity of the shrimp Exhippolysmata oplophoroides (Decapoda: Caridea) in southeastern Brazil

We studied the shrimp Exhippolysmata oplophoroides in southeastern Brazil (Macaé, 22º37″S; Ubatuba, 23º55″S; and Cananéia, 25º53″S) to test the hypotheses that (1) females should have the same reproductive output and fecundity at the regional scale, and (2) these reproductive parameters can vary among regions. Maximum carapace length (Macaé = 13.1 mm, Ubatuba = 12.8, Cananéia = 16), fecundity, and reproductive output differed among regions (Ancova, p < 0.05). Females in Cananéia exhibited the highest reproductive output/fecundity (18.6%/5230 embryos), followed by those in Macaé (16.0%/4365 embryos) and Ubatuba (12.6%/1206 embryos). We found that fecundity/reproductive output varies inversely with the length of the reproductive season, i.e. a considerable amount of energy is allocated during seasonal spawning periods in subtropical regions such as Cananéia. Previous studies have proved that E. oplophoroides is able to reproduce continuously in Ubatuba, with probably minor but continuous energy investment in reproduction occurring in this tropical region. In contrast, although Macaé is located at tropical latitude, it is under the influence of Cabo Frio upwelling that alters the water’s physical properties via cold water intrusion. This process may be a determining factor in the spawning seasonality of Macaé’s shrimp population, justifying the higher reproductive output recorded here than in Ubatuba.

Energy budget adjustment of sea cucumber Apostichopus japonicus during breeding period

Reproductive success is unpredictable for many cultured echinoderms. And cost of reproduction plays an important role in animal life-history. Therefore, understanding cost of reproduction contributes to improving breeding techniques during broodstock rearing in aquaculture. In this study, energetic costs during breeding were explored in the sea cucumber, Apostichopus japonicus, an important aquaculture species in China, Japan and South Korea. The food intake, digestive functions and energy budget patterns were estimated at different breeding stages. Unexpectedly, in the growing gonad phase of the sea cucumber lifecycle, animals showed a decline in feed consumption (from 4.49 to 2.6 g ind−1 day−1), ingestion rate (from 0.4 to 0.13 g g−1 day−1) and apparent digestive rate (from 14.73% to 10.92%), resulting in a reduced energy input. In addition, the increased energy investment in reproduction (from 1.76% to 15.61%) resulted in a decrease in energy allocated to soma growth (from 17.1% to 5.64%) and self-maintenance (from 47.82% to 34.67%). These results suggest that reproduction impairs energy acquisition ability in breeding A. japonicus. And broodstock can adapt energy shortage by internal adjustment of energy allocation strategy.

Geographic variation in life‐history traits: factors shaping energy investment in reproduction in a highland dwelling rodent

AbstractVariation in life‐history traits is shaped by intrinsic and extrinsic factors. Climate influences the availability of energy and nutrients in the environment, and thus animals’ energy balance, their allocation of resources to reproduction and ultimately, fitness. Temperature is reported as a prevailing influence on animals’ investment in reproduction, and diverse hypotheses propose mechanisms by which it occurs. Central and external limits to energy acquisition have been traditionally assumed. Hence, digestive limitations, periods of low food availability or demanding cold conditions would restrict this input. However, under warm conditions endotherms also face the need to dissipate heat from metabolic processes. From this perspective, a high metabolic rate process, such as lactation, would be restricted under increasing ambient temperatures (Ta). Our research addresses the variation in reproductive parameters in Phyllotis xanthopygus, a rodent species broadly distributed in the west highlands and southern parts of South America. Across a pronounced elevation gradient in the central Andes Mountains we found that litter size is comparatively smaller in pregnant females collected at lower elevations. This is congruent with expectations from the heat dissipation limit hypothesis. To disentangle the role of temperature on this pattern, we bred individuals under two thermal treatments. We recorded no changes in the litter size, but pups’ body mass was lower at higher Ta during lactation, also in agreement with decreased metabolism. Offspring size and number are relevant to a species’ fitness. The evidence suggests that in P. xanthopygus these traits vary geographically and are shaped by Ta in the laboratory. A species that evolved in high altitudes, inhabits cold and temperate areas and is sensitive to Ta increases in the laboratory, appears as an excellent candidate to further explore the ecophysiology‐thermal landscape interface, crucial to develop accurate predictive models of biodiversity dynamics.

Demographic responses to warming: reproductive maturity and sex influence vulnerability in an octocoral

Ocean warming, caused by climate change, is critically impacting marine coastal ecosystems. Benthic organisms, such as anthozoans, are increasingly submitted to high temperatures that cause massive mortalities in tropical and temperate seas. To broaden our understanding of their response to thermal stress, we tested the putative role of reproductive maturity and sex in the susceptibility of the Mediterranean red gorgonian, Paramuricea clavata, to high temperatures. We experimentally compared the response to thermal stress of sexually immature (i.e., juveniles) versus mature individuals (i.e., adults), and of males versus females. Colonies' response was firstly assessed by measuring the percentage of tissue area exhibiting necrosis. Then, the reproductive output (i.e., fertility, size, and number of gonads) of both sexes was characterized. When compared to juveniles, adults showed a significantly higher percentage of necrosis, suggesting that during the reproductive period they are more vulner- able to high temperatures. Males and females showed a similar percentage of tissue damage and a significant decrease in their reproductive output. However, females' reproduction was more impacted, suggesting that females are more susceptible to thermal stress than males. A dif- ferential energy investment in reproduction may be the underlying cause of the observed responses. Adults invest a large proportion of their energy budget in reproduction; hence, they have fewer resources available to cope with stress, compared to juveniles. A similar situation seems to apply to females, with respect to males. Considering the current ocean-warming trend, our results imply that the long-term viability of shallow populations of long-lived anthozoans may be jeopardized in the future. This study reveals potential demographic consequences of warming that go beyond its associated increment of mortality rates. Given the important ecological role of many anthozoan species, these results can help better predict the future effects of climate change on coastal ecosystems.

Perchlorate exposure does not modulate temporal variation of whole-body thyroid and androgen hormone content in threespine stickleback

Previously we showed that exposure of threespine stickleback (Gasterosteus aculeatus) to the endocrine disruptor perchlorate results in pronounced structural changes in thyroid and gonad, while surprisingly, whole-body thyroid hormone concentrations remain unaffected. To test for hormone titer variations on a finer scale, we evaluated the interactive effects of time (diel and reproductive season) and perchlorate exposure on whole-body contents of triiodothyronine (T3), thyroxine (T4), and 11-ketotestosterone (11-KT) in captive stickleback. Adult stickleback were exposed to 100ppm perchlorate or control water and sampled at 4-h intervals across the 24-hday and at one time-point (1100h) weekly across the reproductive season (May–July). Neither whole-body T3 nor T4 concentration significantly differed across the day in control or perchlorate treated stickleback. Across the reproductive season, whole-body T3 concentration remained stable while T4 significantly increased. However, neither hormone concentration was significantly affected by perchlorate, verifying our previous studies. The concentration of whole-body 11-KT, a major fish androgen, displayed significant diel variation and also steadily declined across the reproductive season in untreated males; perchlorate exposure did not influence the concentration of 11-KT in either diel or reproductive season schedules. Diel and reproductive season variations in 11-KT content in male stickleback are likely related to reproductive physiology and behavior. The observed increase in T4 content across the reproductive season may be reflective of increased energy investment in reproduction near the end of the life cycle.

The impact of domestication on fearfulness: A comparison of tonic immobility reactions in wild and domesticated finches

We examined differences in the fear response between Bengalese finches and their wild ancestor, the white-backed munia, to explore the evolutionary mechanisms of behavioural changes due to domestication. The Bengalese finch (Lonchura striata var. domestica) was domesticated from the wild-living white-backed munia (L. striata) approximately 250 years ago. A previous study indicated that Bengalese finches sing much more complex songs than white-backed munias. We hypothesised that Bengalese finches are likely able to allocate more resources to reproduction in exchange for reduced survival effort. We measured tonic immobility (TI) reactions as a response to physical restraint to evaluate fearfulness related to coping with predation. The results showed that Bengalese finches exhibited decreased TI responses compared with white-backed munias. TI responses were unaffected by sex, body weight or growth conditions. These differences suggest that the fearfulness in Bengalese finches has been reduced by selective pressure during domestication. Bengalese finches may have been able to increase the investment of energy in reproduction in exchange for reduced costs of predation and coping necessary to survive in the wild; these behavioural changes may have been a major target of domestication effects in this species.

Composition of body storage compounds influences egg quality and reproductive investment in the land snail Cornu aspersum

In invertebrates, resources available for growth and reproduction might influence the composition of body stores and subsequently nutrient allocation to eggs, thereby adjusting energy investment in reproduction. We investigated in the land snail Cornu aspersum (Müller, 1774) the efficiency of growth and the main storage compounds in the body and in eggs with respect to lipid content in food (5.5% versus 2.5%). The high body dry mass density of snails fed on lipid-rich diet underlined the high storage capacity of neutral lipids acquired during growth (high growth efficiency) without changing energy content because of the prevailing carbohydrate storage compounds. Reproductive investment was lower in these snails, and maternal effects decreased clutch size. Triglyceride allocation to eggs might enhance survival probability of offspring and therefore compensate for smaller clutch size. Snails fed on lipid-poor diet maximized their investment in clutch size whatever the amount of body stores, and allocated a higher amount of cholesterol to eggs. Cholesterol could be essential for embryo growth, as it ensures membrane functioning. In conclusion, the availability of resources can differentially affect nutrient allocation and energy investment in reproduction. Thus, the investigation of physiological processes becomes essential to understand population dynamics in fluctuating or changing habitats.

An investigation of the calcification response of the scleractinian coral &amp;lt;i&amp;gt;Astrangia&amp;lt;/i&amp;gt; &amp;lt;i&amp;gt;poculata&amp;lt;/i&amp;gt; to elevated &amp;lt;i&amp;gt;p&amp;lt;/i&amp;gt;CO&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; and the effects of nutrients, zooxanthellae and gender

Abstract. The effects of nutrients and pCO2 on zooxanthellate and azooxanthellate colonies of the temperate scleractinian coral Astrangia poculata (Ellis and Solander, 1786) were investigated at two different temperatures (16 °C and 24 °C). Corals exposed to elevated pCO2 tended to have lower relative calcification rates, as estimated from changes in buoyant weights. Experimental nutrient enrichments had no significant effect nor did there appear to be any interaction between pCO2 and nutrients. Elevated pCO2 appeared to have a similar effect on coral calcification whether zooxanthellae were present or absent at 16 °C. However, at 24 °C, the interpretation of the results is complicated by a significant interaction between gender and pCO2 for spawning corals. At 16 °C, gamete release was not observed, and no gender differences in calcification rates were observed – female and male corals showed similar reductions in calcification rates in response to elevated CO2 (15% and 19% respectively). Corals grown at 24 °C spawned repeatedly and male and female corals exhibited two different growth rate patterns – female corals grown at 24 °C and exposed to CO2 had calcification rates 39% lower than females grown at ambient CO2, while males showed a non-significant decline of 5% under elevated CO2. The increased sensitivity of females to elevated pCO2 may reflect a greater investment of energy in reproduction (egg production) relative to males (sperm production). These results suggest that both gender and spawning are important factors in determining the sensitivity of corals to ocean acidification, and considering these factors in future research may be critical to predicting how the population structures of marine calcifiers will change in response to ocean acidification.

Effects of androgenic gland ablation on growth and reproductive parameters of Cherax quadricarinatus males (Parastacidae, Decapoda)

This work investigates the effects of androgenic gland (AG) ablation on the structure of the reproductive system, development of secondary sexual characters and somatic growth in Cherax quadricarinatus males. The AG ablation, which was performed at an early developmental stage (initial weight: 1.85 ± 0.03 g), had no effect on the somatic growth parameters (specific growth rate and growth increment), but it prevented the re-formation of male gonopores and appendices masculinae. However, the red patch differentiation and chelae size were similar to those in control males. All the ablated animals developed a male reproductive system. Testis structure was macroscopically and histologically normal. The distal portion of the vas deferens (DVD) was enlarged in some animals, with histological alterations of the epithelium and the structure of the spermatophore. Results suggest that the higher growth in males than in females may be due to an indirect effect of the AG on energy investment in reproduction rather than to a direct effect of an androgen. This is the first report of a potential action of the AG on the secretory activity of the distal VD and the structural organization of the spermatophore. Although the AG may play a role in the development of male copulatory organs, its association with the red patch development deserves further research. The results obtained in the present study support and complement those from intersexes of the same species.

The influence of temperature and presence of predators on growth, survival and energy allocation for reproduction in the Catarina scallop Argopecten ventricosus

Environmental factors are known to modify the life history of marine ectotherms. In a 16-month laboratory experiment, we investigated the influence of temperature and presence of predators on life-history parameters including shell growth, survival and the energy investment in reproduction and body mass of the short-lived (2–3 years) scallop Argopecten ventricosus. In parallel, A. ventricosus was maintained in the field at the Pacific coast of Baja California, México, to compare growth, survival and reproductive effort under natural conditions. For the laboratory treatments, scallops were reared at simulated field temperatures (SFT), 5 °C above SFT and in the presence of predators. Elevated water temperatures caused higher growth and gonad production, although at the cost of increased mortality. Presence of predators induced energy allocation to muscle rather than gonad growth, deferred spawning and extended survival. Field scallops exhibited higher growth, higher reproductive investment and were able to reproduce twice, whereas all laboratory scallops died after the first spawning. The natural variability of environmental parameters such as food and temperature may thus support optimal growth in the field, and when animals are protected from predators, reproduction in the second year of life.

Influence of food supply on the δ13C signature of mollusc shells: implications for palaeoenvironmental reconstitutions

Compared to oxygen isotopes, the carbon isotope composition of biogenic carbonates is less commonly used as proxy for palaeoenvironmental reconstructions because shell δ13C is derived from both dissolved inorganic (seawater) and organic carbon sources (food), and interactions between these two pools make it difficult to unambiguously identify any independent effect of either. The main purpose of this study was to demonstrate any direct impact of variable food supply on bivalve shell δ13C signatures, using low/high rations of a 13C-light mixed algal diet fed to 14-month-old (adult) cultured Japanese Crassostrea gigas under otherwise essentially identical in vitro conditions during 3 summer months (May, June and July 2003, seawater temperature means at 16, 18 and 20 °C respectively) in experimental tanks at the Argenton laboratory along the Brittany Atlantic coast of France. At a daily ration of 12% (versus 4%) oyster dry weight, the newly grown part of the shells (hinge region) showed significantly lower δ13C values, by 3.5‰ (high ration: mean of −5.8 ± 1.1‰, n = 10; low ration: mean of −2.3 ± 0.7‰, n = 6; ANOVA Scheffe’s test, p < 0.0001). This can be explained by an enhanced metabolic activity at higher food supply, raising 13C-depleted respiratory CO2 in the extrapallial cavity. Based on these δ13C values and data extracted from the literature, and assuming no carbon isotope fractionation between food and shell, the proportion of shell metabolic carbon would be 26 ± 7 and 5 ± 5% for the high- and low-ration C. gigas shells respectively; with carbon isotope fractionation (arguably more realistic), the corresponding values would be 69 ± 14 and 24 ± 9%. Both groups of cultured shells exhibited lower δ13C values than did wild oysters from Marennes-Ol eron Bay in the study region, which is not inconsistent with an independent influence of diet type. Although there was no significant difference between the two food regimes in terms of δ18O shell values (means of 0.1 ± 0.3 and 0.4 ± 0.2‰ at high and low rations respectively, non-significant Scheffe’s test), a positive δ13C vs. δ18O relationship recorded at high rations supports the interpretation of a progressive temperature-mediated rise in metabolic activity fuelled by higher food supply (in this case reflecting increased energy investment in reproduction), in terms not only of δ13C (metabolic signal) but also of δ18O (seawater temperature signal). Overall, whole-shell δ18O trends faithfully recorded summer/winter variations in seawater temperature experienced by the 17-month-old cultured oysters.

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.