Evolution and consequences of endothermy in fishes

Regional endothermy, the conservation of metabolic heat by vascular countercurrent heat exchangers to elevate the temperature of the slow-twitch locomotor muscle, eyes and brain, or viscera, has evolved independently among several fish lineages, including lamnid sharks, billfishes, and tunas. All are large, active, pelagic species with high energy demands that undertake long-distance migrations and move vertically within the water column, thereby encountering a range of water temperatures. After summarizing the occurrence of endothermy among fishes, the evidence for two hypothesized advantages of endothermy in fishes, thermal niche expansion and enhancement of aerobic swimming performance, is analyzed using phylogenetic comparisons between endothermic fishes and their ectothermic relatives. Thermal niche expansion is supported by mapping endothermic characters onto phylogenies and by combining information about the thermal niche of extant species, the fossil record, and paleoceanographic conditions during the time that endothermic fishes radiated. However, it is difficult to show that endothermy was required for niche expansion, and adaptations other than endothermy are necessary for repeated diving below the thermocline. Although the convergent evolution of the ability to elevate slow-twitch, oxidative locomotor muscle temperatures suggests a selective advantage for that trait, comparisons of tunas and their ectothermic sister species (mackerels and bonitos) provide no direct support of the hypothesis that endothermy results in increased aerobic swimming speeds, slow-oxidative muscle power, or energetic efficiency. Endothermy is associated with higher standard metabolic rates, which may result from high aerobic capacities required by these high-performance fishes to conduct many aerobic activities simultaneously. A high standard metabolic rate indicates that the benefits of endothermy may be offset by significant energetic costs.     

Review: Analysis of the evolutionary convergence for high performance swimming in lamnid sharks and tunas.

Elasmobranchs and bony fishes have evolved independently for more than 400 million years. However, two Recent groups, the lamnid sharks (Family Lamnidae) and tunas (Family Scombridae), display remarkable similarities in features related to swimming performance. Traits separating these two groups from other fishes include a higher degree of body streamlining, a shift in the position of the aerobic, red, locomotor muscle that powers sustained swimming to a more anterior location in the body and nearer to the vertebral column, the capacity to conserve metabolic heat (i.e. regional endothermy), an increased gill surface area with a decreased blood-water barrier thickness, a higher maximum blood oxygen carrying capacity, and greater muscle aerobic and anaerobic enzyme activities at in vivo temperatures. The suite of morphological, physiological, and biochemical specializations that define "high-performance fishes" have been extensively characterized in the tunas. This review examines the convergent features of lamnid sharks and tunas in order to gain insight into the extent that comparable environmental selection pressures have led to the independent origin of similar suites of functional characteristics in these two distinctly different taxa. We propose that, despite differences between teleost and elasmobranch fishes, lamnid sharks and tunas have evolved morphological and physiological specializations that enhance their swimming performance relative to other sharks and most other high performance pelagic fishes.     

The plankton of the tropical western Indian ocean as a biomass indirectly supporting surface tunas (yellowfin,Thunnus albacares and skipjack,Katsuwonus pelamis)

Synopsis The biomass of available forage is a key factor in controlling the abundance and distribution of surface tropical tunas, as they have high energy demands and live in a poor environment. The direct estimate of this forage biomass is not possible with existing techniques. Thus we have investigated the lower link, i.e. the plankton organisms which are the food of fishes preyed upon by tunas. In a previous study, this fraction of the zooplankton has been identified, both by taxa and by size, by analysing the stomach contents of the fishes which are the preys of tunas. In this paper, we use 331 plankton samples from tuna fishing grounds of the tropical Indian ocean, to define the characteristics of the planktonic fraction actually participating in the tuna food chain. Main results are as follows: (1) Only 15–27 % of the total zooplanktonic biomass (> 1 mm) is actually accessible for the fishes preyed upon by surface tunas. This useful part of the zooplankton is a well defined fraction of the planktonic population which remains in the 0–170 meters water layer during daylight hours. This part of the zooplankton accounts for a variable percentage of its total biomass the different geographic areas and represents the most relevant parameter to assess the potential richness of a given area for surface tunas. (2) From areas where fishing for surface tunas is poor to those where fishing is successful, it is observed that the total zooplankton biomass increases by a factor of 4 whereas the biomass of the useful fraction increases by a factor of 7. This disproportionate increase is due to the facts that the potential preys of fishes preyed upon by tunas represent a growing fraction of the zooplankton and that a growing proportion of this fraction remains by day in the 0–170 meter water layer, therefore becoming available for the day-feeders which comprise most of the prey-fishes of surface tunas.     

Endothermy in fishes: a phylogenetic analysis of constraints,predispositions, and selection pressures

Synopsis Endothermy, the ability to raise body temperature by internal heat production, is unusual in teleost fishes and has only been documented within one suborder, the Scombroidei. Two separate modes of endothermy have evolved in the scombroidei; tunas warm their muscles, brain and viscera using heat exchangers in the circulation to these metabolically active tissues while billfishes and one primitive mackerel have a thermogenic organ situated beneath the brain. Both modes of endothermy emphasize common themes. Large body size coupled with heat exchangers are necessary to reduce convective and conductive heat exchange. A tissue with a high oxidative capacity is required for heat generation. Studies based upon morphology and mitochondrial DNA analyses indicate that endothermy has evolved independently at least three times within the scombroid lineage. Mapping of-morphological and physiological traits on a molecular phylogeny for scombroids provides evidence of selective pressures favoring evolution of diverse endothermic styles. The new results suggest anatomical constraints prevent most fish from using the tuna form of endothermy and indicate a possible linkage between endothermy and locomotory style (thunniform or sub-carangiform).Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

Selective advantages conferred by the high performance physiology of tunas,billfishes, and dolphin fish

Tunas are extensively distributed throughout world's oceans and grow and reproduce fast enough to support one of the world's largest commercial fisheries. Yet they are apex predators living in the energy depauperate pelagic environment. It is often presumed that tunas evolved their specialized anatomy, physiology, and biochemistry to be capable of (a) high maximum swimming speeds, (b) high sustained swimming speeds, and/or (c) very efficient swimming, all of which help account for their wide distribution and reproductive success. However, a growing body of data on the energetics and physiological abilities of tunas do not support these assumptions. The three things demonstratively “high performance” about tunas, and probably other pelagic species such as marlin (Makaira spp. and Tetrapturus spp.) and dolphin fish (Coryphaena spp.), are (a) rates of somatic and gonadal growth, (b) rates of digestion, (c) rates of recovery from exhaustive exercise (i.e., clearance of muscle lactate and the concomitant acid load). All of these are energy consuming processes requiring rates of oxygen and substrate delivery above those needed by the swimming muscles for sustained propulsion and for other routine metabolic activities. I hypothesize that the ability of high performance pelagic species (tunas, billfishes, and dolphin fish) to deliver oxygen and metabolic substrates to the tissues at high rates evolved to permit rapid somatic and gonadal growth, rapid digestion, and rapid recovery from exhaustive exercise (abilities central to success in the pelagic environment), not exceptionally high sustained swimming speeds.     

Cardiovascular and respiratory physiology of tuna: adaptations for support of exceptionally high metabolic rates

Synopsis Both physical and physiological modifications to the oxygen transport system promote high metabolic performance of tuna. The large surface area of the gills and thin blood-water barrier means that O₂ utilization is high (30–50%) even when ram ventilation approaches 101 min^–1kg^–1. The heart is extremely large and generates peak blood pressures in the range of 70–100 mmHg at frequencies of 1–5 Hz. The blood O₂ capacity approaches 16 ml dl^–1 and a large Bohr coefficient (–0.83 to –1.17) ensures adequate loading and unloading of O₂ from the well buffered blood (20.9 slykes). Tuna muscles have aerobic oxidation rates that are 3–5 times higher than in other teleosts and extremely high glycolytic capacity (150 mol g^–1 lactate generated) due to enhanced concentration of glycolytic enzymes. Gill resistance in tuna is high and may be more than 50% of total peripheral resistance so that dorsal aortic pressure is similar to that in other active fishes such as salmon or trout. An O₂ delivery/demand model predicts the maximum sustained swimming speed of small yellowfin and skipjack tuna is 5.6 BL s^–1 and 3.5 BL sec^–1, respectively. The surplus O₂ delivery capacity at lower swimming speeds allows tuna to repay large oxygen debts while swimming at 2–2.5 BL s^–1. Maximum oxygen consumption (7–9 × above the standard metabolic rate) at maximum exercise is provided by approximately 2 × increases in each of heart rate, stroke volume, and arterial-venous O₂ content difference.Paper from International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

The aerobic capacity of tunas: Adaptation for multiple metabolic demands

Tunas are pelagic, continuous swimmers, with numerous specializations for achieving a high aerobic scope. Tunas must maintain a high rate of energy turnover, and therefore require elevated levels of aerobic performance in multiple physiological functions simultaneously. Based on a model of oxygen demand and delivery to the swimming musculature, the yellowfin's total oxygen consumption at the predicted maximum sustainable (aerobic) swimming velocity is well below estimates of its maximum oxygen consumption. This suggests that the high aerobic scope of tunas may be a specialization that permits continuous swimming in addition to supplying oxygen to other metabolic functions. Estimates of the metabolic costs of oxygen-debt repayment, growth, and specific dynamic action have been combined with this model of aerobic swimming performance to evaluate the total energy budget in relation to the aerobic scope of the yellowfin tuna. Repayment of the oxygen debt incurred during burst swimming is potentially a large component of tuna respiratory metabolism and the relatively high aerobic capacity of tuna white muscle may be a specialization for rapid lactate clearance.     

Gill morphometrics in relation to gas transfer and ram ventilation in high‐energy demand teleosts: Scombrids and billfishes

This comparative study of the gill morphometrics in scombrids (tunas, bonitos, and mackerels) and billfishes (marlins, swordfish) examines features of gill design related to high rates of gas transfer and the high‐pressure branchial flow associated with fast, continuous swimming. Tunas have the largest relative gill surface areas of any fish group, and although the gill areas of non‐tuna scombrids and billfishes are smaller than those of tunas, they are also disproportionally larger than those of most other teleosts. The morphometric features contributing to the large gill surface areas of these high‐energy demand teleosts include: 1) a relative increase in the number and length of gill filaments that have, 2) a high lamellar frequency (i.e., the number of lamellae per length of filament), and 3) lamellae that are long and low in profile (height), which allows a greater number of filaments to be tightly packed into the branchial cavity. Augmentation of gill area through these morphometric changes represents a departure from the general mechanism of area enhancement utilized by most teleosts, which lengthen filaments and increase the size of the lamellae. The gill design of scombrids and billfishes reflects the combined requirements for ram ventilation and elevated energetic demands. The high lamellar frequencies and long lamellae increase branchial resistance to water flow which slows and streamlines the ram ventilatory stream. In general, scombrid and billfish gill surface areas correlate with metabolic requirements and this character may serve to predict the energetic demands of fish species for which direct measurement is not possible. The branching of the gill filaments documented for the swordfish in this study appears to increase its gill surface area above that of other billfishes and may allow it to penetrate oxygen‐poor waters at depth. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Physiological thermoregulation in bigeye tuna,Thunnus obesus

Synopsis Although a growing body of evidence has indicated that tuna can thermoregulate and have body temperatures that are decoupled from immediate changes in ambient temperature, demonstrating the extent and time-course of body temperature changes in tuna moving through their natural environments has proved to be elusive. Here we use body temperature data telemetered from free-ranging fish to demonstrate short-latency physiological thermoregulation in bigeye tuna. We used a recently developed modeling system to determine the magnitude and time-course of the whole-body thermal conductivity changes that would result in the body temperature changes observed in fish in the wild. The results indicate rapid, 100 to 1000-fold changes in whole-body thermal conductivity that occur in response to quickly changing ambient temperatures. Coupling this physiological response with behavioral thermoregulation expands the foraging space of these animals by permitting activity in wide ranges of water temperatures and depths.Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

Basic concepts relevant to heat transfer in fishes,and their use in measuring the physiological thermoregulatory abilities of tunas

Synopsis Aerobic heat production and heat loss via the gills are inexorably linked in all water breathing teleosts except tunas. These processes are decoupled in tunas by the presence of vascular counter-current heat exchangers, and sustained (i.e., steady state) muscle temperatures may exceed water temperature by 10° C or more in larger individuals. The presence of vascular counter-current heat exchangers is not clearly advantageous in all situations, however. Mathematical models predict that tunas could overheat during strenuous activity unless the efficacy of vascular heat exchangers can be reduced, and that they may be activity limited in warmer waters. Tunas may likewise be forced out of potentially usable habitats as they grow because they have to occupy cooler waters. Vascular counter-current heat exchangers also slow rates of heating and cooling. A reduced rate of muscle temperature decrease is clearly advantageous when diving into colder water to chase prey or avoid predators. A reduced rate of heat gain from the environment would be disadvantageous, however, when fish return to the warmer surface waters. When subjected to changes in ambient temperature, tunas cannot defend a specific body temperature and do not thermoregulate in the mammalian sense. Yet when appropriately analyzed, data taken under steady state and non-steady state conditions indicate that tunas are not strictly prisoners of their own thermoconserving mechanisms. They apparently can modify overall efficiency of their vascular counter-current heat exchangers and thus avoid overheating during bouts of strenuous activity, retard cooling after diving into colder water, and rapidly warm their muscles after voluntarily entering warmer water. The exact physiological mechanisms employed remain to be elucidated.Paper from the International Union of Biological Societies symposium The biology of tunas and billfishes: an examination of life on the knife edge, organized by Richard W. Brill and Kim N. Holland.     

Do method and species lifestyle affect measures of maximum metabolic rate in fishes?

The rate at which active animals can expend energy is limited by their maximum aerobic metabolic rate (MMR). Two methods are commonly used to estimate MMR as oxygen uptake in fishes, namely during prolonged swimming or immediately following brief exhaustive exercise, but it is unclear whether they return different estimates of MMR or whether their effectiveness for estimating MMR varies among species with different lifestyles. A broad comparative analysis of MMR data from 121 fish species revealed little evidence of different results between the two methods, either for fishes in general or for species of benthic, benthopelagic or pelagic lifestyles.     

The standard metabolic rate of dolphin fish

The standard metabolic rates (SMRs) of 11 (1.395–4.125 kg) dolphin fish (mahimahi or dorado, Coryphaena hippurus ) were measured at 25°± 0.5°C. Fish were prevented from swimming with neuromuscular blocking agents and force ventilated. Heart rates were determined simultaneously. SMRs (358–726 mg O₂ h ^–1) were several times those of other similarly sized active teleosts such as salmonids, but close to those of tunas. Heart rates (84–161 beats min ^–1) were also high, but alike those of tunas under similar circumstances. As in tunas, the high SMR of dolphin fish may result from high osmoregulatory costs engendered by their large gill surface areas and/or other adaptations necessary for achieving exceptionally high maximum metabolic rates.     

Complex behaviour by coral-reef fish larvae in open-water and near-reef pelagic environments

We present the first in situ observations of the pelagic larvae of coral-reef fishes feeding, schooling and being preyed upon. In addition, we report on their behavioural interactions with adult and juvenile fishes. Observations on over 500 larvae of over 50 species (mostly from four families) near the end of their pelagic interval were made in both open water (> 1 km offshore) and near-reef environments. Nearly 10% of larvae were seen to feed in open water, but < 1% fed near the reef. Presettlement schooling was observed in five species of four families. We observed no predation upon larvae in open water except near the bottom. Near the reef, 8.5% of larvae were eaten. The main predators near and on the reef were a species of wrasse and lizardfishes. Rates of predation seem to differ among genera of pomacentrids, perhaps related to differences in behaviour when settling. When confronted with adult fishes, which happened largely near the reef, larvae reacted with a limited range of behaviours, including sheltering near the observer, swimming to the surface, slowing or stopping, or swimming offshore. The frequency of these behaviours differed among larvae of three pomacentrid genera. Interactions with reef residents, particularly pomacentrids, were common, and usually involved aggression by the resident toward settling larvae. This may act to discourage settlement during the day when such residents are active. These data show that behaviour of late larvae of coral-reef fishes is complex and can greatly influence survival and recruitment. Further, behaviour differs among taxa, showing that not only are larvae not passive, but also that a generalised behaviour of larvae does not exist.     

Kinetics of growth and sugar consumption in yeasts

An overview is presented of the steady- and transient state kinetics of growth and formation of metabolic byproducts in yeasts.Saccharomyces cerevisiae is strongly inclined to perform alcoholic fermentation. Even under fully aerobic conditions, ethanol is produced by this yeast when sugars are present in excess. This so-called Crabtree effect probably results from a multiplicity of factors, including the mode of sugar transport and the regulation of enzyme activities involved in respiration and alcoholic fermentation. The Crabtree effect inS. cerevisiae is not caused by an intrinsic inability to adjust its respiratory activity to high glycolytic fluxes. Under certain cultivation conditions, for example during growth in the presence of weak organic acids, very high respiration rates can be achieved by this yeast.S. cerevisiae is an exceptional yeast since, in contrast to most other species that are able to perform alcoholic fermentation, it can grow under strictly anaerobic conditions.Non-Saccharomyces yeasts require a growth-limiting supply of oxygen (i.e. oxygen-limited growth conditions) to trigger alcoholic fermentation. However, complete absence of oxygen results in cessation of growth and therefore, ultimately, of alcoholic fermentation. Since it is very difficult to reproducibly achieve the right oxygen dosage in large-scale fermentations, non-Saccharomyces yeasts are therefore not suitable for large-scale alcoholic fermentation of sugar-containing waste streams. In these yeasts, alcoholic fermentation is also dependent on the type of sugar. For example, the facultatively fermentative yeastCandida utilis does not ferment maltose, not even under oxygen-limited growth conditions, although this disaccharide supports rapid oxidative growth.     

Time and tide wait for no fish: intertidal fishes out of water

Synopsis Hypoxic conditions are rare in the open ocean, but may occur during low tides in tidepools. Intertidal fishes respond to low tides in a variety of ways, including avoiding the intertidal zone during low tides, respiring in the well-oxygenated layer at the surface of the water, or simply tolerating hypoxic water. A number of intertidal fish species have the ability to leave the water and survive terrestrially for a period of time while breathing air. This paper reviews the literature on ecomorphology of amphibious intertidal fishes, suggests ecomorphological and ecophysiological approaches to clarifying the adaptations of intertidal fishes for emergence from water, and considers differences in the types of emergence behavior and activities seen in three broadly defined behavioral types. These types include the skippers, fishes that actively emerge at all phases of the tidal cycle and engage in routine terrestrial activity, the remainers, that emerge passively under cover such as rocks or vegetation by remaining in place as the tide recedes, and the tidepool emergers, that typically spend low tides in tidepools but may emerge from hypoxic water. Portioning of gas exchange between the gills and the skin, the release of CO₂ into air, the effect of emergence on metabolic rate, and vertical zonation in distribution of fishes in the intertidal zone are compared for fishes in each of these behavioral styles.     

Scombroid Fishes Provide Novel Insights into the Trait/Rate Associations of Molecular Evolution

The study of which life history traits primarily affect molecular evolutionary rates is often confounded by the covariance of these traits. Scombroid fishes (billfishes, tunas, barracudas, and their relatives) are unusual in that their mass-specific metabolic rate is positively associated with body size. This study exploits this atypical pattern of trait variation, which allows for direct tests of whether mass-specific metabolic rate or body size is the more important factor of molecular evolutionary rates. We inferred a phylogeny for scombroids from a supermatrix of molecular and morphological characters and used new phylogenetic comparative approaches to assess the associations of body size and mass-specific metabolic rate with substitution rate. As predicted by the body size hypothesis, there is a negative correlation between body size and substitution rate. However, unexpectedly, we also find a negative association between mass-specific metabolic and substitution rates. These relationships are supported by analyses of the total molecular data, separate mitochondrial and nuclear genes, and individual loci, and they are robust to phylogenetic uncertainty. The molecular evolutionary rates of scombroids are primarily tied to body size. This study demonstrates that groups with novel patterns of trait variation can be particularly informative for identifying which life history traits are the primary factors of molecular evolutionary rates.     

Modulation of swimming behavior in the medicinal leech

Expression of swimming in the medicinal leech (Hirudo medicinalis) is modulated by serotonin, a naturally occurring neurohormone. Exogenous application of serotonin engenders spontaneous swimming activity in nerve-cord preparations. We examined whether this activity is due to enhanced participation of swim motor neurons (MNs) in generating the swimming rhythm. We found that depolarizing current injections into MNs during fictive swimming are more effective in shifting cycle phase in nerve cords following serotonin exposure. In such preparations, the dynamics of membrane potential excursions following current injection into neuronal somata are substantially altered. We observed: 1) a delayed outward rectification (relaxation) during depolarizing current injection, most marked in inhibitory MNs; and 2) in excitor MNs, an enhancement of postinhibitory rebound (PIR) and afterhyperpolarizing potentials (AHPs) following hyperpolarizing and depolarizing current pulses, respectively. In contrast, we found little alteration in MN properties in leech nerve cords depleted of amines. We propose that enhanced expression of swimming activity in leeches exposed to elevated serotonin is due, partly, to enhancement of relaxation, PIR and AHP in MNs. We believe that as a consequence of alterations in cellular properties and synaptic interactions (subsequent paper) by serotonin, MNs are reconfigured to more effectively participate in generating and expressing the leech swimming rhythm.Abbreviations AHP Afterhyperpolarizing potential - DCC Discontinuous current clamp - DE Dorsal excitor motor neuron - DI Dorsal inhibitor motor neuron - IPSP Inhibitory postsynaptic potential - MN Motor neuron - PIR Postinhibitory rebound - VE Ventral excitor motor neuron - VI Ventral inhibitor motor neuron     

Structural adaptations for ram ventilation: Gill fusions in scombrids and billfishes

For ram‐gill ventilators such as tunas and mackerels (family Scombridae) and billfishes (families Istiophoridae, Xiphiidae), fusions binding the gill lamellae and filaments prevent gill deformation by a fast and continuous ventilatory stream. This study examines the gills from 28 scombrid and seven billfish species in order to determine how factors such as body size, swimming speed, and the degree of dependence upon ram ventilation influence the site of occurrence and type of fusions. In the family Scombridae there is a progressive increase in the reliance on ram ventilation that correlates with the elaboration of gill fusions. This ranges from mackerels (tribe Scombrini), which only utilize ram ventilation at fast cruising speeds and lack gill fusions, to tunas (tribe Thunnini) of the genus Thunnus, which are obligate ram ventilators and have two distinct fusion types (one binding the gill lamellae and a second connecting the gill filaments). The billfishes appear to have independently evolved gill fusions that rival those of tunas in terms of structural complexity. Examination of a wide range of body sizes for some scombrids and billfishes shows that gill fusions begin to develop at lengths as small as 2.0 cm fork length. In addition to securing the spatial configuration of the gill sieve, gill fusions also appear to increase branchial resistance to slow the high‐speed current produced by ram ventilation to distribute flow evenly and optimally to the respiratory exchange surfaces. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Locomotor performance and muscle metabolic capacities: impact of temperature and energetic status

In aquatic ectotherms, muscle metabolic capacities are strongly influenced by exogenous factors, principally temperature and food availability. Seasonal changes in temperature lead many organisms to modify their metabolic machinery so as to maintain capacity even in "slower" cold habitats. Modifications of mitochondrial capacities are central in this response. The increases in protein-specific oxidative capacities of mitochondria during cold acclimation of temperate fishes do not occur during the evolutionary adaptation to cold in Antarctic species. Instead, Antarctic fishes tend to increase the proportion of fibre volume devoted to mitochondria, perhaps to facilitate intracellular distribution of oxygen and metabolites. Variation in energetic status can drastically modify muscle metabolic status, with glycolytic muscle changing more than oxidative muscle. This in turn impacts swimming performance. A decrease in the condition of cod leads endurance at speeds above Ucrit to drop by 70%. Sprint swimming is less affected, perhaps as it does not exhaust glycolytic muscle. We used interindividual variation in muscle metabolic capacities to identify correlates of swimming performance in stickleback and cod. Activities of cytochrome c oxidase in glycolytic muscle are a correlate of sprint swimming in stickleback (Gasterosteus aculeatus) and cod (Gadus morhua), whereas lactate dehydrogenase activities in glycolytic muscle are a correlate of cod endurance swimming. In scallops, gonadal maturation leads to virtually complete mobilisation of glycogen from muscle. This does not reduce the capacity of the scallops, Chlamys islandica and Euvola ziczac, to mount escape responses, but significantly slows their recuperation from exhaustive exercise. Muscle metabolic capacities fall in parallel with glycogen mobilisation. In the compromise between muscles' dual roles as a motor and a macromolecular reserve, a significant loss in locomotory ability occurs during gametogenesis and spawning. Reproductive fitness takes the upper hand over maintenance of performance.     

New expanded bed adsorbents for the recovery of DNA

A 20–40 m pellicular high density (3.7 g cm^–3) expanded bed material has been designed for the capture of DNA and other large macromolecules. Anion exchangers fashioned out of these supports exhibited dramatically enhanced DNA binding capacities over commercial anion exchange adsorbents (6 mg ml^–1, c.f. 50 g ml^–1 at 10% breakthrough), due to a combination of small particle and fuzzy surface architecture created through the coupling of polyethylene imine chains.