A comparison of transport-related gill enzyme activities and tissue-specific free amino acid concentrations of Baltic Sea (brackish water) and freshwater threespine sticklebacks, Gasterosteus aculeatus, after salinity and temperature acclimation

The osmoregulatory abilities of one freshwater and two brackish water (Baltic Sea) populations of the euryhaline teleost fish Gasterosteus aculeatus were studied with respect to evolutionary physiology. Plasma osmolality, activities of Na⁺K⁺-ATPase, citrate synthase, creatine kinase in the gill and free amino acids in liver, axial muscle and pectoral fin muscle were measured. After transfer from 10 to 35 ppt at 15 °C, time-course changes of plasma osmolality and gill Na⁺K⁺-ATPase showed no significant fundamental differences between the freshwater and one of the Baltic Sea populations. In a multi-factorial experiment, each population was exposed to four different abiotic regimes. Both brackish water populations had high mortality in freshwater at 4 °C, which is discussed as a failure of osmotic regulation (reduced taurine concentrations). Freshwater specimens had higher levels of glycine in the axial and pectoral fin muscles compared to the brackish water populations. This is interpreted as a genetically based effect. In brackish (20 ppt) water of 15 °C, the freshwater population had high activities of Na⁺K⁺-ATPase, but low activities of creatine kinase, whereas both brackish water populations behaved in the opposite way. A fundamental difference between the freshwater and brackish water populations on the level of the osmoregulatory machinery was not observed. Received: 10 December 1998 / Accepted: 22 September 1999     

Metabolic enzymes in coelomic cells (eleocytes) of the polychaete Nereis virens: sex specific changes during sexual maturation

The activities of some enzymes of the intermediary metabolism and the content of soluble protein and carbohydrate (glycogen plus free glucose) were measured in one type of coelomic cells (eleocytes) of the polychaete Nereis virens. Specimens used in this study were collected between 1989 and 1991 in Oosterscheldt Bay, The Netherlands, and divided into six different stages of sexual maturation as determined by the mean oocyte volume. In both sexes, the soluble protein content in eleocytes of immature individuals (11 mg ml^–1 cell vol) increased three-fold. In prespawning N. virens the soluble protein content decreased to less than 2 mg protein ml^–1 cell vol in females but not in males. In both sexes, the carbohydrate content decreased continuously from immature [300 mol glucose equivalent (equiv) ml^–1 cell vol] to prespawning individuals (< 40 mol glucose equiv ml^–1 cell vol). During the time course of maturation, the specific activities (expressed as units mg^–1 protein) of pyruvate kinase, phosphoenolpyruvate carboxykinase, malate dehydrogenase, alanine aminotransferase and glutamate dehydrogenase decreased in both sexes. A transient increase in the specific activities was found for glycogen phosphorylase and aspartate aminotransferase. No major changes were found for hexokinase, lactase dehydrogenase, glucose-6-phosphate dehydrogenase and malic enzyme. Sex specific differences were found for the activities of citrate synthase and isocitrate dehydrogenase, which were higher in males. the specific activities of the latter enzyme increased more than ten-fold in males, but only four-fold in female eleocytes during maturation. In eleocytes of prespawning females, the activities of most enzymes showed extremely high variations not found in prespawning males. For two enzymes of fatty acid catabolism, -hydroxyacyl-CoA dehydrogenase and -hydroxybutyrate dehydrogenase, only traces of activities were detected, suggesting the absence of significant fatty acid catabolism in the eleocytes. Compared to the eleocytes, the body wall tissue showed ten-fold higher activities of phosphofructokinase, whereas the eleocytes displayed higher activities of the amino acid interconverting enzymes glutamate dehydrogenase and alanine aminotransferase and the glyconeogenic enzyme phosphoenolpyruvate carboxykinase. Citrate synthase activities were similar for both tissues. In the coelomic fluid of N. virens, glucose (< 0.1 to 3.5 mM) and d-lactate (0.1 to 4 mM) were present and represent exogenous substrates for the eleocyte metabolism.     

Larval mortality during export to the sea in the fiddler crab Uca minax

Dense populations of the fiddler crab Uca minax (Le Conte 1855) are common along tidally influenced freshwater rivers and streams >50 km from the sea. Adults do not migrate from inland sites to release larvae, but instead release them directly into an environment where the zoeae cannot survive. Laboratory salinity tolerance experiments were used to determine how long larvae from the inland-most population of U. minax along the Pee Dee River, South Carolina, USA can survive zero salinity compared to larvae from a brackish water population (salinity 5) near the mouth of Winyah Bay in the same estuary. Larvae from the brackish water population were also exposed to a salinity of 5 and their survival tracked. These experiments were conducted from May to August 2004 and 2005. To determine if inland larvae suffered significant mortality in transit due to salinity stress, current profiles were measured in the field and used to model the time taken by a larva using ebb-tide transport to travel to permissive salinities. The laboratory tolerance experiments showed that larvae from the inland freshwater population had LT50’s of 4–5 days at 0 salinity, which were significantly longer than those of the brackish water zoeae (2–3 days). Zoeae from the brackish water population survived for at least one larval molt at a salinity of 5 with LT 50’s of ∼12 days. Estimated travel times to reach permissive salinities from the inland-most population based on current profiles were 3–5 days for larvae using night-time only ebb-tide transport and 1.5–2.5 days for those using ebb-tide transport both day and night. Previously published field data indicate that U. minax larvae do use both day- and night-time ebb-tide transport, and are found in high densities in the water column during the day. These results lead to the conclusion that U. minax stage I zoeae do not undergo significant salinity-induced mortality during their 50+ km trip to the sea.     

Enzyme activities and allozyme polymorphism in two genetic types (or sibling species) of the genus Marenzelleria (Polychaeta: Spionidae) in Europe

Two populations of Marenzelleria spp. from the North Sea and the Baltic Sea that had already been characterized by allozyme electrophoresis and found genetically different in 1994 were again examined in 1997. The genetic identity (I = 1.0) examined in 1994 and 1997 shows that the structures of the populations of genetic Type I in the North Sea (Ringkobing Fjord) and genetic Type II in the Baltic Sea (Darss–Zingst Bodden chain) are stable. Three diagnostic gene loci that were determined additionally (citrate synthase, one locus; alanine aminotransferase, two loci) support the hypothesis of the existence of two sibling species of the genus Marenzelleria in Europe. In addition, the activities of ten electrophoretically separated enzyme systems were measured in the worms immediately after removal from their habitat and after they had been kept in the laboratory for 2 months. Under both conditions four enzymes showed significant differences between genetic types. The higher activity of the alanine aminotransferase of the North Sea worms (Type I) is discussed in the context of the different preferred salinities of the two Marenzelleria sibling species. Received: 21 January 1999 / Accepted: 13 July 1999     

Activities of metabolic enzymes in the deep-water crabsChaceon fenneri andC. quinquedens and the shallow-water crabCallinectes sapidus

The activities of 18 enzymes were measured in gill, hepatopancreas and muscle tissue of the deep-water crabsChaceon fenneri andC. quinquedens and the shallow-water crabCallinectes sapidus collected from the Gulf of Mexico in January 1989. The activities of catabolic enzymes were correlated in general with the known metabolic rates of the three species. Activities were much higher inC. sapidus than inChaceon fenneri andC. quinquedens. In some cases,C. quinquedens had higher activities thanC. fenneri. The activities of enzymes of amino acid metabolism (glutamate dehydrogenase and alanine aminotransferase) were higher inC. quinquedens, which had high hemolymph [ammonia] and ammonia excretion rates. The activity of lactate dehydrogenase (LDH) ofC. fenneri andC. quinquedens were correlated with the two species' abilities to withstand hypoxia. The more hypoxiatolerant species,C. quinquedens, had higher activity of LDH in its muscles than didC. fenneri.     

An investigation on the anaerobic metabolism of Nephtys hombergii (Annelida: Polychaeta)

In Nephtys hombergii Savigny (Annelida: Polychaeta: Errantia), succinate and less pronounced L-alanine are accumulated as endproducts during an initial phase of anaerobiosis. In this phase aspartate is utilized as substrate for anaerobic energy production in addition to glycogen. Prolonged anaerobiosis results in the formation of propionate and acetate which, to a large extent, are excreted into the water. The concentrations of aspartate and succinate were found to remain unchanged down to a Pw_{O 2}(oxygen partial pressure in incubation water) of approximately 20 torr, indicating a fully aerobic metabolism. At a Pw_{O 2}of 15 torr characteristic changes can be observed: energy production becomes partially anaerobic. At a Pw_{O 2}of 7 torr the level of aspartate is largely reduced, but the accumulation of succinate is still significantly less than at anoxia. [In the habitat of N. hombergii, the intertidal mud flats, a Pw_{O 2}in the range from 0 torr (Pamatmat, 1968) to 12 torr (Wells and Warren, 1975) was measured.] The activities of some important enzymes involved in anaerobic energy production (among them pyruvate kinase, phosphoenolpyruvate carboxykinase and lactate dehydrogenase) were measured. Although N. hombergii has a high glycolytic capacity, no lactate dehydrogenase was detected. Instead strombine and alanopine dehydrogenases were present in comparatively high activities.     

Protective effects of honey against cadmium-induced alteration of some biochemical parameters in rats

The objective of this study is to elucidate on the effect of honey on plasma and organ biochemical parameters of albino rats exposed to acute and sub-chronic dose of cadmium chloride. Uric acid levels, activities of plasma lactate dehydrogenase (LDH), aspartate aminotransferase (AST) and alanine aminotransferase (ALT) activities, organ (liver, lungs, and kidney) AST, ALT, superoxide dismutase (SOD), and catalase (CAT) were determined. Plasma uric acid and glucose levels, plasma LDH, AST, and ALT activities were significantly decreased; activities of organ SOD, CAT, AST, and ALT activities were increased in rats administered sub-chronically and acutely with cadmium (Cd). Treatment with honey restored their levels to normal.     

Reproductive investment reduces recuperation from exhaustive escape activity in the tropical scallop Euvola zizac

In scallops, mobilization of reserves from the adductor muscle to support maintenance and reproductive activity may impinge upon a major role of the adductor muscle, the movement of the valves during swimming and escape responses. The tropical scallop Euvola ziczac (Linnaeus 1758) invests energy reserves to different degrees during its two periods of reproduction each year. We evaluated the impact of reproductive investment on the escape responses (clapping capacity and recovery after exhaustive exercise) of E. ziczac sampled at different reproductive stages (immature, mature, spawned) during the two reproductive periods in 1997. We compared the escape response capacities with the levels of muscle energetic reserves (glycogen, proteins, and arginine phosphate) and muscle metabolic capacities [activity of the glycolytic enzymes: glycogen phosphorylase (GP), pyruvate kinase (PK), phosphofructokinase (PFK), octopine dehydrogenase (ODH), arginine kinase (AK), and the mitochondrial enzyme, citrate synthase (CS)]. Gonad size, gamete volume fraction, and levels of gonadal protein and lipid were greater for mature scallops during the first than during the second reproduction. Numbers of claps during escape responses (49–57) and levels of muscle arginine phosphate remained similar throughout the different reproductive stages in both reproductive periods. In contrast, recovery was slowed during gonadal maturation in both reproductive periods and during spawning in the first reproduction. Scallops generally took more time to regain their initial clapping capacity during the first (25–40 min) than during the second reproduction (20–30 min). Muscle glycogen decreased markedly during both gonadal maturation and spawning in both reproductions; whereas, muscle proteins decreased only in the first reproduction. The levels of most enzymes decreased during gonadal maturation in both reproductions, and also after spawning, particularly during the first reproduction. We concluded that gonadal maturation and spawning did not decrease clapping capacity of E. ziczac, but decreased its capacity to recover from exhaustive exercise most likely due to decreased levels of energetic reserves and a reduced metabolic capacity of the adductor muscle. Moreover, these effects were probably stronger during the first cycle because of the greater reproductive investment coincident with decreased food availability. Received: 28 April 2000 / Accepted: 21 September 2000     

Metabolic enzyme activities in shallow- and deep-water chondrichthyans: implications for metabolic and locomotor capacity

Biochemical indices of white (WM) and red muscle (RM) aerobic and anaerobic metabolic capacity were measured in 14 species of benthic and benthopelagic chondrichthyans from a depth of ~90 to 2,200 m to evaluate the relationship between metabolic capacity and depth of occurrence, phylogeny, and locomotor mode. Maximal activities of the enzymes citrate synthase, malate dehydrogenase (MDH), lactate dehydrogenase (LDH), and pyruvate kinase (PK) were analyzed in muscle tissue at 10 °C. These were combined with previously published elasmobranch data in order to represent a comprehensive range of depths, phylogeny, and locomotor modes (i.e., benthic, benthopelagic, pelagic). Significant decreases in WM PK and LDH activities and a lack of significant trends in RM enzyme activities with increasing median depth of occurrence (MDO) indicate a depth-related reduction in both burst-locomotor and metabolic capacity. These trends are consistent with the “visual-interactions hypothesis.” Phylogeny and locomotor mode had little influence on enzyme activities compared to MDO, and the present study suggests similar activities in co-occurring demersal sharks and rays. Overall, the present study indicates low metabolic capacities in deep-sea chondrichthyans, which is important to consider when managing deep-sea fisheries.     

Haemolymph constituent levels and ammonia efflux rates of Nephrops norvegicus during emersion

Nephropsnorvegicus (L.) were subjected to 8 h of emersion, either between layers of seawater-soaked hessian with periodical (20 min) flushes of seawater (high humidity, HH) or to unprotected emersion (low humidity, LH). Blood ammonia levels rose during emersion in both groups but reached higher levels under LH conditions. Ammonia efflux rates after re-immersion were higher than those of control prawns, and amounts of ammonia excreted at such times were considerably higher than those calculated to have accumulated in the blood during emersion. Possible explanations for such differences are discussed. C _aO2 and C _vO2 decreased rapidly to ca. 10% normoxia values within 2 h of HH and LH emersion and remained low throughout the remaining emersion time. Emersion-induced tissue hypoxia increased blood concentrations of glucose and lactate. Lactate accumulation was higher during LH emersion, compared with HH emersion. Blood pH dropped ca. 0.40 units but increased again after 2 h of re-immersion. Acidosis was probably related more to respiratory difficulties (CO₂ accumulation) than to lactate accumulation, as blood lactate values remained high after 2 h of re-immersion. The ability of N. norvegicus to cope with emersion appears to be little influenced by high humidity conditions. Received: 26 June 1996 / Accepted: 5 August 1996     

Gender-based comparative toxicity of di-ethyl phthalate in Wistar rats

In recent years, the exposure of humans to phthalate esters through environmental contamination has increased. One among them is di-ethyl phthalate (DEP), which is used as a plastisizer for cellulose ester plastic films and sheets, solid rocket propellants, molded and extruded articles, as a component in insecticide sprays and various other substances, as well as in industrial applications. Release into the environment occurs primarily as a result of production and manufacturing of DEP and during the use and disposal of products containing DEP. Therefore, a study was undertaken to evaluate gender-specific toxicity of DEP in Wistar rats. Rats of both sexes, weighing 125–130?g, were administered 50?ppm (w/v) DEP in water ad libitum for a period of 180 days and were given normal diet. Control animals received normal diet and water ad libitum. During the treatment, rats were weighed every week and water consumption per day was measured. After the completion of treatment, liver weight?:?body weight^?1 ratio, liver weight, body weight^?1, liver and serum enzymes, and other biochemical parameters of liver and serum were assessed. It was observed that there was no significant change in body weight^?1, liver weight, liver weight?: body weight^?1 ratio, and water consumption in both sexes. There were significant increases in liver acid phosphatase (ACP) activity and kidney glutathione levels, and nonsignificant changes in liver alkaline phosphatase (ALP), aspartate aminotransferase (AST), alanine aminotransferase (ALT), succinate dehydrogenase (SDH), and lactate dehydrogenase (LDH) activities in DEP-treated male rats, whereas in DEP-treated female rats the liver showed significant decrease in ALP and SDH and nonsignificant changes in AST, ALT, and LDH activities. Serum ACP and LDH levels in DEP-treated male rats were significantly decreased, and in the case of DEP-treated female rats, only serum LDH levels were significantly decreased. There was no significant change in serum ALP, AST, and SDH levels in DEP-treated male and female rats as compared to control rats. Histology of the livers of both male and female rats showed loss of hepatic architecture, degenerative changes in hepatocytes, and vacuolation in hepatocytes in both the centrilobular and periportal areas. It can be concluded from this study that prolonged exposure to DEP at 50?ppm levels can be harmful to animals and humans. This is evident from the present study as certain significant changes in enzyme activities in the liver, serum, and histological alterations in liver were observed.     

Occurrence of marine resident tropical eel Anguilla bicolor bicolor in Indonesia

In order to understand the migratory history and habitat use of the tropical anguillid eels Anguilla celebesensis, A. marmorata, and A. bicolor bicolor, the otolith strontium (Sr) and calcium (Ca) concentrations were examined for eels collected in Indonesian waters. In A. bicolor bicolor collected in a lagoon, the change in Sr:Ca ratios outside the high Sr:Ca core generally indicated two patterns of habitat residence: (1) constant living in either brackish or sea waters with no freshwater life (25%) and (2) habitat shifts from fresh water to brackish or sea waters (75%). No A. bicolor bicolor had a general life history as a freshwater resident. A. celebesensis and A. marmorata from the uppermost freshwater lake showed freshwater life history patterns. The wide range of otolith Sr:Ca ratios in A. bicolor bicolor indicated that the habitat use of this tropical eel was facultative among fresh, brackish, and marine waters during the growth phase after recruitment to coastal areas similar to that for temperate eels. Thus, the migration of anguillid eels into fresh water is clearly not an obligatory.     

Sex-biased dispersal in a long-lived polygynous reptile (Crocodylus johnstoni )

We compared natal dispersals of freshwater crocodiles (Crocodylus johnstoni) against the prediction of male dispersal bias for a polygynous mating system. The crocodiles inhabited a linear series of pools and we calculated the net distances from natal pools to recapture locations some 12–18 years later, at maturity. Philopatry was assessed in terms of adult social distances. A female social distance was 0.46 pools and a male social distance was 1.0 pool. By these criteria, both sexes showed low levels of philopatry (7–12%). However, individuals of both sexes dispersed from the natal site long before they were sexually mature. Divergence in dispersal patterns by sex occurred after the maturity threshold, as males dispersed two to three times farther than females. Intrasexual competition by males is resolved by a size-based hierarchy. The displacement of small males from local mating access is a probable cause of the longer dispersals undertaken by males. Competition, rather than inbreeding avoidance, is driving dispersal in this population of freshwater crocodiles. Received: 8 May 1998 / Accepted after revision: 26 June 1998     

L-Lactate dehydrogenase in the longitudinal muscle of the sea cucumber Sclerodactyla briareus (Echinodermata: Holothuroidea)

The infaunal holothurian Sclerodactyla briareus (Thyone briareus) is able to tolerate exposure to hypoxic conditions for over 2 days. Since the in vitro anaerobic degradation of glucose-U-C¹⁴ by longitudinal muscle preparations leads to an accumulation of labeled lactic acid, it is apparent that lactate dehydrogenase plays a key role during anoxia. Disc electrophoresis resolved one major band of lactate dehydrogenase (LDH) activity in longitudinal muscle extracts. Muscle extracts of s. briareus oxidized L-lactate only, as is the case with most other echinoderms. The apparent Michaelis constants for lactate oxidation and pyruvate reduction are dissimilar from values reported for other marine invertebrates. The LDH activity in this holothurian is higher than the activities reported for other marine invertebrates with the exception of some active, large arthropods. Substrate inhibition of LDH maximal activity occurs at substrate concentrations (pyruvate) exceeding 10 mM, and then to only a minor extent. The characteristics of lactate dehydrogenase from the longitudinal muscle of S. briareus support the hypothesis that it functions to maintain glycolytic flux during prolonged periods of exposure to anoxic or hypoxic conditions.     

Male reproductive success: paternity contribution to queens and workers in Formica ants

The relative number of workers and female sexuals fathered by two males mated with a queen were directly assessed using microsatellite and allozyme markers in field colonies of the ants Formica exsecta and F. truncorum. In both species one of the two males consistently fathered more offspring than the other. There was, however, no evidence that one male might be particularly successful in fathering a disproportionally high proportion of female sexuals relative to the proportion of workers. Moreover, in F. exsecta, the proportions of worker pupae and worker adults fathered by each male did not differ significantly between cohorts. The most likely explanation for this pattern is that females store different amounts of sperm from the two males they mated with. Received: 10 January 1997 / Accepted after revision: 22 March 1997     

Effects of nutrition and temperature on metabolic enzyme activities in larval and juvenile red drum,Sciaenops ocellatus,and lane snapper,Lutjanus synagris

The metabolic enzyme activities were determined in larvae of red drum, Sciaenops ocellatus, and lane snapper, Lutjanus synagris, to determine the effect of temperature and nutrition on metabolic enzyme activities and to evaluate if metabolic enzyme activities are useful in assessing the feeding condition of larval fish. During experiments conducted during the spring of 1990, lactate dehydrogenase (LDH) activities in both red drum and lane snapper were approximately an order of magnitude lower than values typical for adult fish; LDH and citrate synthase (CS) activities increased during early developmental stages, but nutritional effects were apparent. Clear differences (up to 4-fold) between well-fed and starving fish were evident in both LDH and CS activity in red drum. Differences between well-fed and poorly fed larvae were evident until 9 d after hatching. Lane snapper larvae reared at a 25°C had significantly lower LDH activities than larvae reared at 28°C.     

Mating system and alternative male mating tactics in the grass goby Zosterisessor ophiocephalus (Teleostei: Gobiidae)

 Peculiar fertilization dynamics, with males releasing sperm in mucous trails lasting several hours, characterize some demersal spawning fish. The mating system was investigated in a natural population of one of these species: the grass goby Zosterisessor ophiocephalus (Pallas, 1814), a large coastal goby inhabiting seagrass meadows in shallow brackish water. Adult males ranged in size from 7.4 to 23 cm total length, but only larger ones were observed to dig and defend a burrow, where they performed parental care on eggs laid by one to several females. Field observations together with analyses of age, sperm production, trail sperm content and sperm competition tests indicated the occurrence of alternative male mating tactics, likely the expression of an ontogenetic gradient. Larger males are older than smaller ones, and while the former are territorial, the latter “sneak” territorial male spawns. The ejaculate characteristics indicate that grass goby males have functionally polymorphic spawns: in fact sperm trails of larger males last longer and release fewer sperm than those of smaller males. Sperm production over several days is more constant in larger than in smaller males, but the total number of sperm released is higher in the latter. The influence of seminal fluid in the functional intraspecific variability in sperm release in this species is discussed. Received: 30 December 1999 / Accepted: 31 July 2000     

Ontogeny of osmoregulation in the European sea bass Dicentrarchus labrax L.

 Under natural conditions, sea bass eggs hatch in the open sea and young-of-the-year sea bass are found close to estuaries and enter brackish and even freshwater lagoons in the Mediterranean and the eastern Atlantic. The ontogeny of osmoregulation in the European sea bass, Dicentrarchus labrax (Linnaeus, 1758), was studied in different developmental stages, from hatching to large juveniles, exposed to a range of salinities at 17 °C. The experiments were carried out from March to August 1998 in southern France. The type of hyper-hypo-osmotic regulation did not change during development. All stages hyper-regulated at low salinity (under 10–11‰) and hypo-regulated at higher salinities. The acquisition of the full ability to hypo-regulate occurred in four steps. Osmoregulatory capacity was size- and age-dependent and reached its maximum for fish 17–26 mm long, 63–86 days after hatching. The iso-osmotic salinity was 10.2–11.6‰. Our results suggest that early development of osmoregulatory ability, and thus of salinity tolerance in sea bass, may provide an advantageous flexibility for the timing of migration towards low-salinity habitats. Received: 22 May 2000 / Accepted: 18 December 2000     

Sex-specific nest defense in house sparrows (Passer domesticus) varies with badge size of males

According to indicator models of sexual selection, females can benefit from choosing males with above average epigamic traits, but empirical evidence for such benefits is scarce. Here, we report results from an experiment with 29 pairs of house sparrows (Passer domesticus) where the intensity of nest defense against a mounted mustelid predator was related to the size of the black throat and breast patch (“badge”) in males. Using principal components analysis (PCA), original response variables of both sexes were reduced to two factors: “Approach” to the predator and “Distant warning”. “Approach”, the more risky behavior, increased from small- through medium- to large-badged males and decreased in their females. Since large-badged males have a higher certainty of paternity (i.e. greater benefits from defense) and may be older and more experienced (i.e. incur lower costs), the most likely explanation for male defense intensity increasing with badge size is an improving benefit/cost ratio. The resulting optimal response of their females and evolutionarily stable participation in joint parental care is illustrated by a graphical model. It shows that females would, indeed, benefit directly from choosing large-badged males. This, however, is no proof of a direct evolutionary tie between badge size and paternal behavior, as assumed by indicator models of sexual selection. It may simply represent a spurious relationship, originating from the correlation of badge size and defense with confidence of paternity. Received: 22 September 1997 / Accepted after revision: 3 November 1997     

Strontium/Calcium ratios in statoliths of the neon flying squid, Ommastrephes bartrami (Cephalopoda), in the North Pacific Ocean

Strontium to calcium ratios were observed along longitudinal sections of statoliths of nine neon flying squid, Ommastrephes bartrami (LeSueur, 1821), including three mature females (422 to 454 mm mantle length, ML; 207 to 306 d old) obtained from the North Pacific (27–35°N; 144–150°E) during winter and six immature males and females (187 to 226 mm ML; 126 to 164 d old) collected from 39°N; 145°E and 39°N; 169°W during summer. The distances between the nucleus (core) and the edge of the dorsal dome were approximately 660 to 690 μm in mature females and 450 to 510 μm in the immature squid. Sr/Ca ratios were determined at intervals of 30 μm between the nucleus and edge of the dorsal dome. Sr/Ca ratios were higher in areas near the nuclei and peripheral portions of the dorsal dome than in the middle portions of the statoliths (270 to 420 μm from the nuclei, corresponding to ages of 60 to 90 d) in mature females; thus a U-shaped pattern was evident. Sr/Ca ratios in the six immature squid decreased from nucleus to the dorsal dome; in three squid the ratios slightly increased toward the dorsal dome edge. The observed Sr/Ca ratios in immature squid were considered to represent younger portions of the U-shaped pattern. In the present study we discuss this pattern in relation to environmental and biological conditions of O. bartrami, which undertakes seasonal migrations between spawning grounds in the Subtropical Domain and feeding grounds in the Subarctic Domain and Transitional Zone in the North Pacific Ocean. Although Sr/Ca ratios are potentially affected by ambient water temperature and ontogenetic conditions, including somatic growth and statolith growth, it was impossible to evaluate each environmental and biological effect separately, as variations in these factors are complicated and effects could be interdependent. Received: 11 April 1997 / Accepted: 27 December 1997