Effects of limiting access to prey on development of first zoeal stage of the brachyuran crabs Cancer magister and Hemigrapsus oregonensis

We tested the influence of limiting access to prey on larval development of the crabs Cancer magister and Hemigrapsus oregonensis by raising their Stage 1 larvae in the laboratory on different prey densities and with various periods of access to prey. Experiments were conducted in 1995 and 1996 at the Shannon Point Marine Center in Anacortes, Washington, USA. Our results show that crab larvae do not require continuous access to prey for optimal development nor do they appear to require light for prey capture. Survival and duration of Stage 1 C. magister fed continuously on only one-fourth the amount of the control density of prey and those fed at the control density for only 6 h per day were the same as for larvae fed continuously at the control density (20 ml⁻¹). Larvae with cyclic access to prey at the control density for 24 h and then starved for 72 h showed significantly lower survival and longer instar duration to Stage 2. Experiments on Stage 1 H. oregonensis which investigated a combination of prey density, period of access to prey and light/dark conditions during feeding revealed that survival decreased with decreasing prey density or with decreasing feeding period, but no differences were observed during periods of limited prey availability as a function of light or dark conditions. Stage duration was not affected by reduced prey density nor by the light/dark condition at the time of feeding, but it was prolonged when the period of access to prey was limited. The period of access to prey did not affect the weight of Day 1 Stage 2 larvae. Larvae fed high densities of prey for 4 h followed by 20 h of reduced-density diet exhibited the same survival and stage duration as controls that were continuously fed high-density prey. Our results define sub-optimal diets that can be used experimentally to determine the nutritional contributions made by naturally-occurring prey organisms during larval development in the two species. In nature, larvae may satisfy nutritional requirements through periodic encounters with dense prey patches during vertical migrations by day or night. Received: 12 August 1997 / Accepted: 5 February 1998     

Experimental investigation into the effects of suspended sediment on fertilisation, larval survival and settlement in a scleractinian coral

Laboratory and field experiments were used to determine whether high (≃100 mg l⁻¹), low (≃50 mg l⁻¹) and control (≃0 mg l⁻¹) levels of suspended sediment affected fertilisation, larval survival, and larval settlement in the scleractinian coral Acropora digitifera (Dana, 1846). Both high- and low-sediment treatments significantly decreased fertilisation, but post-fertilisation embryonic development was not inhibited by suspended sediments. Larval survival and larval settlement were significantly reduced in high- and low-sediment treatments. No difference was found between high- and low-sediment treatments in any of the three post-spawning processes investigated, suggesting that they are susceptible to sediment concentrations which are not exceptionally high even under natural conditions (>50 mg l⁻¹). The introduction of an additional stress in the form of high levels of suspended sediments coupled with naturally high variability in recruitment may have a considerable effect on the successful supply and settlement of coral larvae to a reef. Given that many coral communities are open reproductive systems, the consequences of disturbance events are not likely to be restricted to the impact area. Recruitment to a population may be reduced significantly in the presence of high levels of suspended sediments because of effects on larval survival and settlement. Recruitment of larvae to adjacent populations may also be affected due to a decreased fertilisation success and potential increases in mortality of larvae passing through the affected site. Received: 13 August 1998 / Accepted: 22 July 1999     

Physiological effects of the detergent linear alkylbenzene sulphonate on blue mussel larvae (Mytilus edulis) in laboratory and mesocosm experiments

A series of laboratory (short-term exposure in small beakers) studies and a 19 d mesocosm (6 m³ polyethylene bags filled with fjord water) study were conducted on blue mussel, Mytilus edulis, larvae and plantigrades exposed to a concentration gradient of the detergent linear alkylbenzene sulphonate (LAS, 0 to 39 mg l⁻¹). LAS is increasingly found in nearshore environments receiving wastewater from urban treatment plants. The aims were to observe physiological effects on swimming, grazing and growth in the laboratory and effects on settling and population development at in situ conditions (in field mesocosms) in order to evaluate the damages on ciliated meroplankton caused by LAS. In the laboratory the larvae showed a 50% mortality at 3.8 mg LAS l⁻¹ after 96 h exposure whether or not food was provided. Additionally the swimming behaviour was affected at 0.8 mg LAS l⁻¹ (i.e. a more compact swimming track, a smaller diameter of the swimming tracks, and reduced swimming speed). The larval particle grazing was reduced 50% at 1.4 mg LAS l⁻¹. The specific growth rate of the larvae was reduced to half at 0.82 mg LAS l⁻¹ over 9 d. During the mesocosm experiment, the larval population showed a dramatic decrease in abundance within 2 d at concentrations as low as 0.08 mg LAS l⁻¹, both due to a significantly increased mortality, but also due to settling. The settling success was reduced at the same LAS concentration as that at which mortality was observed to increase significantly. In addition to reduced settling rate, the larvae showed delayed metamorphosis and reduced shell growth as a response to LAS. Our hypothesis that the larval ciliary apparatus, crucial for normal swimming, orientation, and settling behaviours and for particle uptake, was damaged due to LAS exposure is supported by our results. This is confirmed by the physiological data (grazing, growth) and in the direct video-based observations of larval performance (swimming) and provides a reasonable explanation for what was observed in the bags (abundance, settling, mortality). These physiological effects on blue mussel larvae/plantigrades occurred at LAS concentrations reported to occur in estuarine waters. Received: 15 January 1997 / Accepted: 12 February 1997     

Diel variability of feeding activity in haddock (Melanogrammus aeglifinus) larvae in the East Shetland area, North Sea

Investigations of factors affecting feeding success in fish larvae require knowledge of the scales of variability of the feeding process itself and the indices used to assess this variability. In this study, we measured short-term (diel) variability in feeding rates of wild haddock (Melanogrammus aeglifinus) larvae four times per day during a 10-d cruise in the northern North Sea. Feeding activity was evaluated using indices of gut fullness, prey digestive state and biochemical measurements (tryptic enzyme activity). The gut fullness and the enzyme activity indices indicated moderate to high rates of food consumption throughout the cruise. Time series analysis of the three indices showed significant diel variability in all indices and enabled identification of significant lags between food uptake and peak digestive enzyme activity. The typical pattern of food consumption and digestion was characterized by maximal ingestion of prey early in the evening (19:00 hrs) and peak digestive enzyme activity at 01:00  hrs. The time scale over which enzyme activities reacted to prey ingestion was ca. 6 h, and is consistent with expectations from controlled laboratory experiments with other larval fish species. Significant diel variability in tryptic enzyme activity suggests that attempts to relate this measure of feeding success to other variables (e.g. food concentrations) should take care to accommodate natural cycles in feeding activity before making statistical comparisons. Received: 29 October 1998 / Accepted: 18 June 1999     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Feeding ecology and evolutionary survival of the living coelacanth Latimeria chalumnae

One concept of evolutionary ecology holds that a living fossil is the result of past evolutionary events, and is adapted to recent selective forces only if they are similar to the selective forces in the past. We describe the present environment of the living coelacanth Latimeria chalumnae Smith, 1939 at Grande Comore, western Indian Ocean and report depth-dependent cave distribution, temperature, salinity and oxygen values which are compared to the fish's distribution and its physiological demands. We studied the activity pattern, feeding behaviour, prey abundance and hunting success to evaluate possible links between environmental conditions, feeding ecology and evolutionary success of this ancient fish. Transmitter tracking experiments indicate nocturnal activity of the piscivorous predator which hunts between approximately 200 m below the surface to 500 m depth. Fish and prey density were measured between 200 and 400 m, both increase with depth. Feeding tracks and feeding strikes of the coelacanth at various depths were simulated with the help of video and laser techniques. Along a 9447 m video transect a total of 31 potential feeding strikes occurred. Assuming 100% hunting success, medium-sized individuals would obtain 122 g and large females 299 g of prey. Estimates of metabolic rates revealed for females 3.7 ml O₂ kg⁻¹ h⁻¹ and for males 4.5 ml O₂ kg⁻¹ h⁻¹. Today coelacanths are considered to be a specialist deep-water form and to inhabit, with their ancient morphology, a contemporary environment where they compete with advanced, modern fish. Received: 5 July 1999 / Accepted: 11 November 1999     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000     

Characteristics of feeding on dinoflagellates by newly hatched larval crabs

The zoeal larvae of brachyuran crabs must feed soon after hatching on a diet that includes large micro- and mesozooplankton in order to satisfy nutritional requirements. However, newly hatched larvae have been shown to ingest a variety of dinoflagellates, perhaps using microbial carbon sources to sustain them until they encounter more favored prey. Ingestion of dinoflagellates by larval crabs has been documented previously under conditions in which the larvae were exposed to algae provided in monoculture or in defined mixtures of cells. We report here on experiments conducted on the hatching stage of five crab species to determine if ingestion of dinoflagellates occurred when they were provided in combination with Artemia sp. nauplii or after a period of feeding on mesozooplankton. Quantitative measurements of chl a in the larval guts provided evidence of ingestion of algal cells. Active ingestion of the dinoflagellate Prorocentrum micans at specified intervals during an extended feeding period was determined on larvae of two crab species using fluorescently labeled cells provided for brief periods at prescribed time intervals. Stage 1 larvae of four of the five crab species ingested dinoflagellates when they were provided in combination with nauplii and larvae of all five species ingested cells after feeding solely on nauplii for 24 h. Ingestion of algal cells was first evident in the larval guts after 6 h of feeding at both low (200 cell ml⁻¹) and high (1,000 cells ml⁻¹) prey densities. Higher prey densities resulted in higher gut chl a. Larvae continuously exposed to dinoflagellates actively ingested cells at every 3 h interval tested over a 36 h period. Results confirm previous studies that larvae will ingest dinoflagellates even when they are encountered in a mixed prey field or when having previously fed. Ingestion of cells may occur on a continual basis over time.     

Feeding rates of the jellyfish Aurelia aurita on fish larvae

We quantified feeding rates of field caught Aurelia aurita feeding on yolk sac cod (Gadus morhua) larvae in a series of incubation experiments. A short-time (~1 h) functional response experiment with a wide range of prey concentrations (0.5–16 prey l⁻¹, initial concentration) revealed that ingestion rates increased linearly over this range, such that clearance rates were similar between the different prey concentrations. This suggests that A. aurita is capable of efficiently utilizing dense prey patches. This indication was further supported by a linear increase of prey captured by A. aurita during 2.5 h of feeding at extremely high prey concentration (>200 prey l⁻¹). Clearance rate in darkness scaled with jellyfish diameter to a power of ~1.7 for jellyfish 3.9–9.5 cm in diameter. The jellyfish did not alter their umbrella pulse frequency in response to presence of fish larvae. There were no significant differences between A. aurita feeding rates in light and darkness for yolk sac prey ages 0–7 days (at 7.5°C). Although prey vision and escape abilities of fish may develop rapidly during early larval ontogeny, these factors apparently have little impact on interactions with predators such as A. aurita during the yolk sac stage.     

Prey, feeding rates, and asexual reproduction rates of the introduced oligohaline hydrozoan Moerisia lyonsi

Moerisia lyonsi Boulenger (Hydrozoa) medusae and benthic polyps were found at 0 to 5‰ salinity in the Choptank River subestuary of Chesapeake Bay, USA. This species was introduced to the bay at least 30 years before 1996. Medusae and polyps of M. lyonsi are very small and inconspicuous, and may occur widely, but unnoticed, in oligohaline waters of the Chesapeake Bay system and in other estuaries. Medusae consumed copepod nauplii and adults, but not barnacle nauplii, polychaete and ctenophore larvae or tintinnids, in laboratory experiments. Predation rates on copepods by medusae increased with increasing medusa diameter and prey densities. Feeding rates on copepod nauplii were higher than on adults and showed no saturation over the range of prey densities tested (1 to 64 prey l⁻¹). By contrast, predation on copepod adults was maximum (1 copepod medusa⁻¹ h⁻¹) at 32 and 64 copepods l⁻¹. Unexpectedly, M. lyonsi colonized mesocosms at the Horn Point Laboratory during the spring and summer in 4 years (1994 to 1997), and reached extremely high densities (up to 13.6 medusae l⁻¹). Densities of copepod adults and nauplii were low when medusa densities were high, and estimated predation effects suggested that M. lyonsi predation limited copepod populations in the mesocosms. Polyps of M. lyonsi asexually produced both polyp buds and medusae. Rates of asexual reproduction increased with increasing prey availability, from an average total during a 38 d experiment of 9.5 buds polyp⁻¹ when each polyp was fed 1 copepod d⁻¹, to an average total of 146.7 buds polyp⁻¹ when fed 8 copepods d⁻¹. The maximum daily production measured was 8 polyp buds and 22 medusae polyp⁻¹. The colonizing potential of this hydrozoan is great, given the high rates of asexual reproduction, fairly wide salinity tolerance, and existence of a cyst stage. Received: 29 October 1998 / Accepted: 3 March 1999     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Essential fatty acid (docosahexaenoic acid, DHA) availability affects growth of larval herring in the field

Larval fish growth and survival depends not only on prey quantity, but also on prey quality. To investigate effects of prey fatty acid concentration on larval herring growth, we collected different prey organisms and larval herring (Clupea harengus L.) in the Kiel Canal during the spring season of 2009. Along with biotic background data, we analysed fatty acids both in prey organisms and in the larvae and used biochemically derived growth rates of the larvae as the response variable. Larval herring reached their highest RNA/DNA derived growth rates only at high docosahexaenoic acid (DHA) concentration. When the ratio of copepodids to lesser quality cirriped nauplii was low, larval growth and larval DHA concentration were both significantly negatively affected. This was true even as prey abundance was increasing. This finding indicates that even in mixed, natural feeding conditions, growth variations are associated with DHA availability in larval fish.     

Nitrogen balance in marine fish larvae: influence of developmental stage and prey density

The utilization and fate of nitrogen in larvae of plaice (Pleuronectes platessa), blenny (Blennius pavo) and herring (Clupea harengus), from the stage of first-feeding to metamorphosis, was examined under laboratory conditions. Rates of ammonia excretion, primary amine defaecation, and growth in terms of protein-nitrogen were monitored throughout larval life. Data were used to calculate daily ration, the coefficient of nitrogen utilization (absorption efficiency), and gross and net growth efficiencies. The developmental pattern of nitrogen balance was similar for plaice and blenny larvae. These species showed increasing growth efficiency (k₁: 55 to 80%) with decreasing weight-specific waste nitrogen losses with age. Absorption efficiencies. were high (83 to 98%) in plaice and blenny larvae, and tended to increase with development in the former species. Ration relative to body weight decreased with growth in both species. Herring larval development, although at a slower rate than blenny and plaice, appeared normal up to 33 d, after which high mortality occurred. Absorption efficiency in this species tended to decline (83 to 43%) with age, until metabolic costs exceeded the absorbed ration and growth ceased. Artemia sp. nauplii proved a suitable food source for the rearing of plaice and blenny larvae, but this diet may have long-term toxicity or deficiency effects on herring. Availability and density of food affected nitrogen balance in the larvae of all three species. Feeding stimulated the output of wastes in excretion and defaecation by a factor of up to ten times the 12-h non-feeding basal rates. Waste nitrogen output reached a peak some 2 to 3 h after commencement of feeding and returned slowly to the baseline in 5 to 10 h after cessation of feeding. There was an asymptotic increase in ration, ammonia output and growth of larvae as prey density increased. Ration saturated at a higher prey density (>4 prey ml^-1) than either growth or excretion rate (1 prey ml^-1). Thus the efficiency with which food is absorbed and utilized for growth must eventually decline in response to high prey density. The idea that larval fish are adapted to maximize ingestion and growth rate, rather than optimize growth efficiency and thus to respond to prey occurring in either low density or in occasional patches, is supported by these results.     

Size-specific changes in amount of lipid and daily growth rate of early larval sardine, Sardinops melanostictus, in the main Kuroshio Current and its offshore waters off eastern Japan

The size-specific nutritional conditions of larval sardines, Sardinops melanostictus, from the main Kuroshio Current and its offshore waters off eastern Japan were assessed by lipid analysis. A rapid lipid analysis technique (diagnostic kits for human serum lipids) was used to measure the different lipid components of individual sardine larvae as indicators of their nutritional condition. Size-specific growth trajectories of individual larvae were estimated by the biological intercept method, and the recent daily growth rate of standard length (SL) was calculated from the 3 d outer increment width on the otolith. Relationships between the amount of larval phospholipid (PL; tissue weight indicator) and SL, and the recent daily growth rate of larva (Gr) and SL, could be expressed by the equations PL = 0.459 SL ^1.77 and Gr = 0.0809 SL − 0.341, respectively. There was no notable difference in these values between the two survey areas. The relationship between the amount of triglyceride (TG) and SL could be expressed allometrically (TG = 0.013 SL ^2.63). The relationship between the index of starvation tolerance (TG/PL) and SL could be expressed by the allometric equation TG/PL = 0.0288 SL ^0.865, suggesting that larger sardine have a higher starvation tolerance than smaller sardine. The TG of the 8 to 9 mm SL size-class larvae in the offshore area was higher than in the main Kuroshio Current. To test whether the TG for each larval sardine in the 8 to 9 mm SL size-class could be correlated with variables (temperature, chlorophyll a and distribution density of the sardine larvae) measured at the sampling stations, correlation analyses were performed. A highly significant negative correlation between TG and distribution density of the sardine larvae was found. A density-dependence effect seemed to influence the fluctuation of the larval storage energy component for short-term needs. Received: 12 March 1998 / Accepted: 26 March 1999     

Prolonged exposure to low dissolved oxygen affects early development and swimming behaviour in the gastropod Nassarius festivus (Nassariidae)

Effects of low dissolved oxygen on early development and swimming behaviour of veliger larvae of the scavenging gastropod Nassarius festivus were studied. Embryonic development was significantly delayed when dissolved oxygen level was reduced to 3.0 mg O₂ l⁻¹ and no embryo hatched successfully at 0.5 mg O₂ l⁻¹. Veliger larvae hatched at 4.5 mg O₂ l⁻¹ had significantly smaller velar lobe, shell length and shell width. Median 48-h LC₅₀ value of the veliger larvae was estimated at 1.25 mg O₂ l⁻¹ with lower swimming speed (swimming velocity and dispersal velocity) being recorded for the survivors exposed to reduced oxygen levels. The percentage of veliger larvae that developed into crawling juveniles was significantly reduced and metamorphosis was delayed at 4.5 mg O₂ l⁻¹ whereas all larvae at 3.5 mg O₂ l⁻¹ died before they underwent metamorphosis. Juveniles developed at 4.5 mg O₂ l⁻¹ were also smaller than those at 6.0 mg O₂ l⁻¹. Results indicated that dissolved oxygen levels well above hypoxia levels (2.8 mg O₂ l⁻¹) have already had significant impact on the hatching success and larval development in gastropods, which may lead to long-term decreases in population growth.     

Asymptotic growth trajectories of larval sardine (Sardinops melanostictus ) in the coastal waters off western Japan

Growth trajectories of individual larvae of Japanese sardine, Sardinops melanostictus, caught in the coastal waters off western Japan were back-calculated from the first feeding stage up to date of capture (approximate size of 20 to 35 mm total length; TL) based on individually determined allometric relationships between otolith daily ring radii and fish total lengths. The larvae in January-, February-, and March-hatched cohorts in the coastal waters grew faster and more uniformly than those in the oceanic waters offshore of the Kuroshio current. Growth trajectories of the three hatch-month cohorts were similar and could be expressed by the Gompertz model. The inflection points of the growth curves were reached at 9 to 11 d after hatching, when larvae were 10.8 to 11.8 mm TL. Maximum growth rates at these points were 0.80 to 0.85 mm d⁻¹. Growth rates gradually declined after the inflection points, and larval TLs converged into the infinite length of 29 to 32 mm, the sizes at which metamorphosis from larvae to juveniles is initiated. This asymptotic growth pattern in the larval stage resulted in the narrow ranges in TLs in spite of the wide range of ages of the larvae caught by boat seiners in the coastal waters. Slow growth and therefore long duration of the metamorphosing stage could be influential in determining the cumulative total mortality in the early life stages of the Japanese sardine. Received: 14 July 1996 / Accepted: 20 August 1996     

Variations in the content of trypsin and trysinogen in larval herring (Clupea harengus) digesting copepod nauplii

Trypsin and its proform trypsinogen were quantified by radioimmunoassay in herring (Clupea harengus L.) larvae subjected to different prey densities. During the first weeks of larval life, the enzyme content fluctuated in a threephased pattern. Yolk resorption (Phase 1) was characterized by an increase in enzyme. During the first few days after yolk resorption (Phase 2), there was a sharp decline in enzyme. Older larvae (Phase 3) exhibited a second period of intensive enzyme synthesis. Amounts of trypsin in intestines of feeding larvae were analysed. At first feeding, a basal level of gut enzyme of approximately 30ng was recorded, and the amount of additional enzyme secreted from the pancreatic tissue into the intestine appeared to be dependent upon the numbers of prey items ingested. The enzyme-substrate ratio in the intestine was approximately 1 to 4. Prey availability affected amount of trypsinogen. Larvae experiencing a high prey density had an approximately two-fold higher specific enzyme content in Phase 2 compared to larvae exposed to a low prey density. A proposed nutritional strategy for first feeding herring larvae is discussed.     

Effects of inorganic and organic phosphates on feeding, feeding absorption, nutrient allocation, growth and righting responses of the sea urchin Lytechinus variegatus

The sea urchin Lytechinus variegatus is capable of surviving chronic exposure to sodium phosphate (inorganic phosphate) concentrations as high as 3.2 mg l⁻¹, and triethyl phosphate (organic phosphate) concentrations of 1,000 mg l⁻¹. However, chronic exposure to low (0.8 mg l⁻¹ inorganic and 10 mg l⁻¹ organic phosphate), medium (1.6 mg l⁻¹ inorganic and 100 mg l⁻¹ organic phosphate) or high (3.2 mg l⁻¹ inorganic and 1,000 mg l⁻¹ organic phosphate) sublethal concentrations of these phosphates inhibits feeding, fecal production, nutrient absorption and allocation, growth and righting behavior. Food consumption and fecal production declined significantly in individuals exposed to medium and high concentrations of inorganic phosphates and all levels of organic phosphates. Feeding absorption efficiencies for total organics and carbohydrates decreased significantly in individuals held in the highest concentration of organic phosphate. Feeding absorption efficiencies for lipids were significantly reduced in the highest inorganic phosphate concentration only, while they decreased significantly for protein with increasing phosphate exposure. Carbohydrate and lipid levels in gonad and gut tissues decreased significantly with exposure to increasing phosphate concentrations, potentially impairing both gametogenesis and nutrient storage in the gut. Moreover, gonad indices significantly decreased in individuals exposed to the highest concentrations of either phosphate. Growth rates decreased significantly under the influence of all phosphate concentrations, while increasing in seawater alone. Individuals exposed to increasing phosphate concentrations showed reduced righting responses (a measure of stress) and no acclimation in righting times during chronic exposure to phosphates over a 4 week period. These findings indicate that shallow-water populations of L. variegatus subjected to inorganic and organic phosphate pollutants will exhibit stress and be inhibited in their growth and performance due to reductions in feeding, nutrient absorption and allocation of nutrients to key somatic and reproductive tissues. Received: 10 April 2000 / Accepted: 2 October 2000     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.