Behavioral Responses of Walleye Pollock, Theragra Chalcogramma, Larvae to Experimental Gradients of Sea Water Flow: Implications for Vertical Distribution

Walleye pollock larvae under controlled laboratory conditions were exposed to vertical gradients of sea water flow in low and high light. Whether flow originated from the surface or the bottom, larvae responded by altering depth distribution, showing attraction to low flows, avoidance of higher flows and when flow was above a threshold level, loss of ability to orient, swim and feed. These results demonstrate that walleye pollock have the capability for responding to gradients of flow by adjusting their vertical distribution. Walleye pollock and many other pelagic fish larvae have weak swimming capabilities and are generally unable to directly control horizontal distributions in the sea by swimming in higher flow regimens. However, using vertical migration, larvae may select conditions of flow direction and speed which are favorable for feeding and predator avoidance and which indirectly allow them to control transport, aggregation and dispersion.     

Phototactic responses of unfed walleye pollock,Theragra chalcogramma larvae: comparisons with other measures of condition

Synopsis The capability of unfed walleye pollock, Theragra chalcogramma, larvae to swim horizontally towards light was used as a sensitive, sublethal measure of larval condition. At 9°C, positive phototaxis and swimming ability of larvae was fully developed by 4–6 d after hatching, then decreased steadily until death by 12 d after hatching. This measure of larval condition corresponded closely with previously established benchmarks of larval condition, including first feeding, yolksac absorption, point of no return and death by starvation. The presence and timing of behavioral deficits associated with starvation, such as decreased ability to swim, feed and avoid predators, may have significant effects on the ability of larvae to vertically migrate, avoid predators and find and capture food.     

Effects of size and light on respiration and activity of walleye pollock (Theragra chalcogramma) larvae

The respiration rate and swimming activity of walleye pollock (Theragra chalcogramma) larvae were measured in the laboratory to determine how these were affected by body size (measured as dry weight), and amount of light. Size influenced respiration rates, but not activity. Activity increased with increased light, and as walleye pollock larvae developed, light had an increasingly important effect on respiration rate. For older larvae, light is an important factor affecting respiration rate and this may be due to an increased sensitivity to light. Thus, in addition to size, light plays an important role in the energetics of walleye pollock larvae.     

The effect of early and late hatching on the escape response of walleye pollock (Theragra chalcogramma) larvae

Hatching of fish eggs fertilized at the same time occurs overa period of several days. Differences in the escape responseof fish larvae during the hatching period have not hithertobeen studied. In this study, the escape response of walleyepollock (Theragra chalcogramma) larvae over the hatching periodwas examined. Escape speed, response to multiple touches witha fine probe, response to water currents generated by a predatorand predation by euphausiids (Thysanoessa inermis) and amphipods(Pleusirus secorrus) were measured in the laboratory. Otolithmeasurements of field-collected larvae support a broad hatchingperiod for walleye pollock eggs in the sea similar to that observedin the laboratory. The escape response of walleye pollock larvaewas affected by rank in the order of hatching, thus with respectto predation, hatching order may affect the survival of larvaein the sea. Early hatching larvae were smaller, less sensitiveto tactile stimulation, had a slower, weaker escape responseand higher laboratory rates of predation mortality than thosethat hatched later.     

Effects of Low Salinity on Adult Behavior and Larval Performance in the Intertidal Gastropod Crepipatella peruviana (Calyptraeidae)

Shallow-water coastal areas suffer frequent reductions in salinity due to heavy rains, potentially stressing the organisms found there, particularly the early stages of development (including pelagic larvae). Individual adults and newly hatched larvae of the gastropod Crepipatella peruviana were exposed to different levels of salinity stress (32(control), 25, 20 or 15), to quantify the immediate effects of exposure to low salinities on adult and larval behavior and on the physiological performance of the larvae. For adults we recorded the threshold salinity that initiates brood chamber isolation. For larvae, we measured the impact of reduced salinity on velar surface area, velum activity, swimming velocity, clearance rate (CR), oxygen consumption (OCR), and mortality (LC50); we also documented the impact of salinity discontinuities on the vertical distribution of veliger larvae in the water column. The results indicate that adults will completely isolate themselves from the external environment by clamping firmly against the substrate at salinities ≤24. Moreover, the newly hatched larvae showed increased mortality at lower salinities, while survivors showed decreased velum activity, decreased exposed velum surface area, and decreased mean swimming velocity. The clearance rates and oxygen consumption rates of stressed larvae were significantly lower than those of control individuals. Finally, salinity discontinuities affected the vertical distribution of larvae in the water column. Although adults can protect their embryos from low salinity stress until hatching, salinities <24 clearly affect survival, physiology and behavior in early larval life, which will substantially affect the fitness of the species under declining ambient salinities.     

The influence of light on egg buoyancy and hatching rate of the walleye pollock, Theragra chalcogramma

Changes in buoyancy in fertilized bathypelagic eggs of the walleye pollock, Theragra chalcogramma , collected from Shelikof Strait in the Gulf of Alaska were measured under controlled laboratory conditions in density gradient columns from 90 h post–fertilization through hatching. Eggs were incubated at 6° C and exposed to either diel light or constant dark. Eggs held under diel light conditions became more dense than eggs under constant dark beginning <10 h after exposure to light and remained so until 12 h before hatching. Eggs held under constant dark then became more dense than those under diel light. Hatching of eggs under both conditions began at the same time but eggs under diel light showed a delayed hatching rate. Light–induced changes in egg density indicate the ability of walleye pollock eggs to respond to external stimuli and thereby alter their position in the water column in an ecologically meaningful way.     

Quantitative composition,distribution, and feeding of large jellyfish (<Emphasis Type="Italic">Scyphozoa et Hydrozoa</Emphasis>) on the West Kamchatka shelf in summer

The total biomass of jellyfish on the shelf of the eastern Sea of Okhotsk in the summer is estimated as 1672700 tons according to the results of hydroacoustic measurements and 901000 tons by the method of squares. The use of hydroacoustic technologies makes evaluation of the actual stock and range of medusae more accurate, and the further enhancement and perfection of the hydroacoustic method based on multifrequency measurements enables one to obtain more reliable estimates. A significant increase (nearly 25 times) of the total jellyfish biomass takes place in the summer period. Cyanea prevailed in biomass in the spring and Chrysaora melanaster prevailed in the summer. Some species showed considerable expressed spatial differentiation of distribution and affinity to certain environmental conditions. The studied species were almost exclusively zoophages. Their algal diet consisted mainly of diatom algae. Scyphomedusa’s diet mainly included the so-called “peaceful” zooplankton, viz., euphausiids and copepods (as a rule, over 50% the mass), at the same time carnivorous zooplankton, saggits, amphipods, and small medusae also formed a substantial share of their diet. One individual of the predominant jellyfish species consumes a total of 6.1 to 70.5 kcal during its lifecycle, which corresponds to 79.1–513.0 g of raw organic material, assuming 70% assimilability. The relatively low demand for food of this sort can be explained by the low caloric value of the jellyfish body, 96–97% of which consists of water. The distribution and composition of the jellyfish prey show that scyphomedusae exert the greatest influence on the nekton community, as they concentrate in the shelf area of the eastern part of the sea, at walleye pollock spawning sites. There the larvae of bottom invertebrates, including commercially valuable organisms, such as crab and shrimp, are also consumed. In the summer, jellyfish eat nearly 100 billion eggs and 20 billion larvae of walleye pollock, as well as 130 billion decapod (mostly crab) larvae each day, which corresponds to 0.03% of the eggs and 0.003% of the larvae of walleye pollock and 0.003% of the decapod larvae in the estimated stock.     

Diel patterns of behavior in juvenile walleye pollock,Theragra chalcogramma

Synopsis In laboratory experiments, we contrasted the behavior of juvenile walleye pollock under dark and light conditions, comparing group cohesion, activity levels, vertical distribution, and movement into cold water when the water column was thermally stratified. At night the fish displayed a tendency for movement into the upper water column and were less active, with groups more dispersed than during the day. Time spent in cold water under stratified conditions did not differ between day and night. These results are interpreted relative to the potential ecological benefits and costs of particular behaviors, and their application in designing effective sampling surveys of juvenile abundances.     

Microzooplankton, vertical mixing and advection in a larval fish patch

A large ({small tilde}30 ? 75 km) patch of larval walleye pollock,Theragra chalcogramma, was located south of the Alaska Peninsuladuring May 1986. A drifter deployed in this patch followed ananticyclonic path consistent with dynamic topography. Changesin community composition and vertical distribution of microzooplankton>40 µm were sampled for 4 days alongside this drifterto examine feeding conditions for larvae. Biological and physicalchanges during the first 2 calm days revealed substantial small-scalevariability within the larger circulation pattern. Changes duringthe last 2 days were dominated by vertical mixing due to strongwinds. Despite mixing, prey concentrations remained adequatefor feeding by larval pollock as determined by laboratory studies.A satellite-tracked drifter replaced the first drifter and wasstill located within the patch 6 days later. Overall distributionsof larvae and movements of the drifters show a net translationof 7.8 km day^–1 south-westward, but details of the studyreveal complex interactions between coastal waters and a coastalcurrent. During the 10-day period there was an increase in standardlength of the larval fish population of 0.13 mm day^–1and a decline in abundance of {small tilde}7.6% day^–1.Both calculated rates must be underestimates due to continuingrecruitment of small larvae from hatching eggs.     

Soil pH influences embryonic survival in Pseudophryne bibronii (Anura: Myobatrachidae)

Abstract: There has been limited research examining the role that terrestrial habitat characteristics play in influencing persistence of amphibian populations. In this study we investigated the influence of a terrestrial habitat attribute on the distribution of the terrestrial egg‐laying toadlet, Pseudophryne bibronii. Eggs of this species are deposited in depressions, or under leaf litter, and develop to a stage where they can hatch to free swimming tadpoles when water covers breeding sites or embryos are washed into water. Because rainfall can be intermittent, eggs may sit for extended periods in terrestrial nests before sufficient rainfall initiates hatching. Appropriate egg‐laying sites must be chosen by both males and females to ensure embryos survive these periods. A study of 20 sites found that soil pH at sites where P. bibronii were recorded as present was significantly lower than pH at sites where P. bibronii were recorded as absent. To determine whether soil pH influenced the hatching success in P. bibronii, experiments were conducted in which batches of eggs were translocated to local areas with different soil pH. Survival rate of eggs was significantly influenced by relatively small changes in pH, with hatching success higher at lower pH. In a laboratory trial survival of eggs raised on sterilized soil was not affected by soil pH. Fungal infection of eggs was observed in the field and laboratory suggesting that hatching success might be affected via a complex interaction between soil pH, fungi and other soil biota. Choice trials using P. bibronii metamorphs indicated that individuals were capable of distinguishing between small differences in pH. Therefore, distribution of P. bibronii may be influenced by either differential mortality of embryos, or habitat choice by post metamorphic individuals, and further experiments are required to determine the relative importance of these factors.     

Reproductive conditions of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> (Gadidae) off the northeastern coast of Sakhalin Island,Sea of Okhotsk

The spatial distribution of eggs and larvae of the walleye pollock Theragra chalcogramma is considered in respect to dynamics of oceanologic processes, nutrients, chlorophyll а and zooplankton off the northeastern coast of Sakhalin Island in spring 2012. It is shown that the effect of severe temperature regime in the near-bottom horizons in the western Sea of Okhotsk during the spawning period of walleye pollock becomes milder due to specific features of water dynamics. The egg distribution is determined by mesoscale eddies in the region. The species survival depends on the effect of such environmental factors as freshwater discharge from the Amur River, eddy structure in waters of the Sea of Okhotsk, and dynamics of phytoplankton and zooplankton development.     

The influence of predator presence on utilization of artificial seagrass habitats by juvenile walleye pollock,Theragra chalcogramma

Synopsis Behavioral preference for a structured habitat (artificial seagrass) by juvenile walleye pollock,Theragra chalcogramma, was tested in controlled laboratory experiments. We monitored position of fish in 2000 1 tanks with and without artificial seagrass present in one half of the tank. In addition, we exposed walleye pollock to a predator model, assessing their response when a grass plot was available or unavailable as a potential refuge. In the absence of predators, the fish avoided the artificial seagrass, displaying a preference for the open water side of the experimental tanks. In the presence of a predator model, however, juvenile walleye pollock readily entered the artificial seagrass plots. In addition, they often remained in the grass canopy in proximity to the predator instead of moving out of the grass to avoid the predator (when no grass was present they consistently moved to the opposite side of the tank from the predator). The behavioral choices exhibited in this study suggest that juvenile walleye pollock modify habitat selection in response to perceived predation risk, and recognize the structure provided by artificial seagrass as a potential refuge.     

Chemoreception and vertical movement in planktonic yolk-sac larvae of red sea bream Pagrus major

The capability of planktonic yolk-sac larvae of red sea bream Pagrus major in detecting food was examined in the laboratory to ascertain basic knowledge on the early life history of this marine fish. After infrequent vertical burst swimming followed by slight rising or sinking, the larvae remained motionless within thin layers of concentrated food extract (rotifer, Brachionus plicatilis). At the moment of hatching, the larvae already have receptor cells with several cilia arranged radially in their open nostrils. Thus it is likely that by means of their vertical movement they are capable of sensing the thin food patch layer. We suggest that planktonic larvae of Pagrus major are capable of detecting and remaining within food patches even before the onset of feeding. The onset of food detection in the earlier stages may be, to some extent, the more efficient strategy for larval survival and growth because this ability could contribute to a reduction in energy consumption.     

Embryonic development of the bullseye puffer Sphoeroides annulatus (Tetraodontidae): A morphofunctional approach to ontogenetic steps

The embryonic development of the bullseye puffer, Sphoeroides annulatus, was characterized on the basis of the theory of saltatory ontogeny. This theory predicts a correlative relationship between the ontogeny-type in an altricial-precocial spectrum and the habitat that a species occupies within an unstable-stable environmental spectrum. Because S. annulatus inhabits a variety of unstable environments along a wide latitudinal range, the hypothesis that this species presents one of the most altricial embryonic developments among tetraodontids was tested. Based on major developmental events that marked the ontogenetic thresholds nine embryonic steps were identified. Developmental features such as small adhesives eggs, lack of vitelline circulation, small free embryos swimming up at hatching guided by positive phototaxis, and small first-feeding larvae actively swam in the water column, suggest that S. annulatus belongs to the reproductive guild of the nonguarders-lithopelagophils. Moreover, a comparative analysis of the developmental sequences, egg size, and first-feeding larvae size between tetraodontids confirms the hypothesis of this study and supports the evolutionary principle of the altricial-precocial spectrum postulated in the theory of saltatory ontogeny.     

Social and reproductive behavior of a captive group of walleye pollock,Theragra chalcogramma

Synopsis The social and reproductive behavior of a group of four male and seven female walleye pollock,Theragra chalcogramma, were observed in a large tank. Pollock spent most of their time swimming in a loose aggregation near the surface. Males descended from the aggregation more often than females to follow and make physical contact with other males as well as with females. The difference between males and females in the frequency of diving in our tank is consistent with the reported pattern of depth segregation of the sexes in natural pollock spawning aggregations. The frequency of social interactions increased when pollock became reproductively active and was higher at night and during twilight when most of the spawning occurred. Male interactions with females most frequently involved physical contact, while male interactions with other males were more often limited to following. There was no indication that male-male interactions result in the formation of stable social dominance relationships that determine priority of access to mates, as has been suggested previously for walleye pollock. Rather, following and contact interactions appear to promote male identification of potential mates and encounters with ripe females. The possible functional significance of male social interactions is discussed in relation to reports on natural walleye pollock spawning aggregations.     

Behavioral responses to food odor in juvenile marine fish: Acuity varies with species and fish length

This study tested the hypothesis that acuity of behavioral responses to food odor in three commercially important species of marine fish would increase as juvenile length increased. Swimming activity among two size groups of fish was measured in the presence of a series of squid extract dilutions. Increased swimming activity in juvenile Pacific halibut Hippoglossus stenolepis Schmidt, walleye pollock Theragra chalcogramma Pallas, and sablefish Anoplopoma fimbria Pallas was stimulated above threshold concentrations of squid extract, expressed as dilution from full strength. Maximum chemosensory acuity was observed in smaller (8-14 cm total length, TL) Pacific halibut and walleye pollock, while larger sablefish (15-23 cm TL) continued to develop acuity. Response thresholds were highest (10^− 3 dilution) in Pacific halibut, at intermediate levels (10^− 4-10^− 6 dilution) in walleye pollock and smaller sablefish and reached the lowest levels (10^− 13 dilution) in larger sablefish. The widely held view that dissolved free amino acids (DFAA) are the primary chemosensory stimulants for fish food searching may not be valid for sablefish, as they detected squid extract at dilutions containing DFAA that appeared to be far below ambient sea water DFAA concentrations.     

Formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma,larvae under laboratory conditions: role of visual and chemical stimuli

Synopsis Although planktonic marine fish larvae are often distributed in aggregations, the role of behavioral responses to environmental factors in these aggregations is not well understood. This work examines, under laboratory conditions, the influence of visual and chemical stimuli in the formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma, larvae. Larvae were exposed to a horizontal gradient of light (visual stimulus), prey scent (chemical stimuli: squid/copepod and rotifer) or prey density (visual & chemical stimuli: rotifers). While larvae did not respond to prey scent, they did respond to a gradient of light or prey, which resulted in the formation and maintenance of aggregations. Larvae moved into and remained in a zone of higher light intensity (0.56 versus 0.01 mol photons m^-2 s^-1). Once encountering a patch of prey, larvae remained aggregated within the patch to feed. In nature, movement of walleye pollock larvae in response to selected environmental factors (e.g., gravity, light, temperature, turbulence) may serendipitously bring them into contact with prey patches, where they then could remain to feed as long as light intensity remained at or above levels necessary for feeding.     

Minute tubercles in bitterling larvae: Developmental dynamic structures to prevent premature ejection by host mussels

1. Bitterlings are small freshwater fish that use long ovipositors to lay eggs in host mussels, and they have morphological adaptations to increase larval survival. The most well‐known adaptation is the minute tubercles on the skin surface of larvae; they are developed in early‐stage larvae with weak swimming ability and disappear in free‐swimming larvae before they leave the host mussel.
2. In the present study, I comprehensively analyzed the developmental stages of Rhodeus pseudosericeus larvae, their morphological and physiological characteristics, their migration inside mussels, and the development of minute tubercle in order to elucidate the morphological function of the minute tubercles. These tubercles began to develop 1 day after hatching (formation stage), grew for 2–5 days (growth stage), reached the peak height after 6–7 days (peak stage), abruptly reduced in height after 8–10 days (abrupt reduction stage), and gradually reduced in height (reduction stage) until completely disappearing 27 days after hatching (disappearance stage).
3. The larvae remained in the mussels’ interlamellar space of the gill demibranchs until 10 days after hatching and began to migrate to the suprabranchial cavity 11 days after hatching. At this time, the larvae had a heart rate and the caudal fin began to develop. At 24 days after hatching, the minute tubercles had almost disappeared, and some individuals were observed swimming out of the mussels.
4. The results presented herein elucidate that the minute tubercles are the developmental dynamic structures that the bitterling larvae have morphologically adapted to prevent premature ejection from the mussel.

     

Swimming Speed of Larval Snail Does Not Correlate with Size and Ciliary Beat Frequency

Many marine invertebrates have planktonic larvae with cilia used for both propulsion and capturing of food particles. Hence, changes in ciliary activity have implications for larval nutrition and ability to navigate the water column, which in turn affect survival and dispersal. Using high-speed high-resolution microvideography, we examined the relationship between swimming speed, velar arrangements, and ciliary beat frequency of freely swimming veliger larvae of the gastropod Crepidula fornicata over the course of larval development. Average swimming speed was greatest 6 days post hatching, suggesting a reduction in swimming speed towards settlement. At a given age, veliger larvae have highly variable speeds (0.8–4 body lengths s⁻¹) that are independent of shell size. Contrary to the hypothesis that an increase in ciliary beat frequency increases work done, and therefore speed, there was no significant correlation between swimming speed and ciliary beat frequency. Instead, there are significant correlations between swimming speed and visible area of the velar lobe, and distance between centroids of velum and larval shell. These observations suggest an alternative hypothesis that, instead of modifying ciliary beat frequency, larval C. fornicata modify swimming through adjustment of velum extension or orientation. The ability to adjust velum position could influence particle capture efficiency and fluid disturbance and help promote survival in the plankton.     

The abundance of Anisakis simplex L3 in the body-cavity and flesh of marine teleosts

Smith J. W. 1984. The abundance of Anisakis simplex L3 in the body-cavity and flesh of marine teleosts. International Journal for Parasitology14: 491–495. In experiments conducted at sea, whole (ungutted) fish of three species were stored on ice at 3–5 °C for periods from 0 to 72 h. Some Anisakis simplex L3 migrated from the body-cavity into the flesh in mackerel Scomber scombrus, but not in either blue whiting Micromesistius poutassou or whiting Merlangius merlangus. Earlier work has shown that larvae migrate into the flesh of herring Clupea harengus post mortem but not of walleye pollock Theragra chalcogramma. It seems, therefore, that larvae migrate post mortem into the flesh of ‘fatty’ species (e.g. herring, mackerel) but not of ‘non-fatty’ species (e.g. blue whiting, whiting, walleye pollock). No significant larval excapsulation occurred in isolated mackerel or whiting viscera. In vivo, most encapsulated L3 occur in the body-cavity of euphausiid-feeding fish (herring, mackerel, blue whiting, walleye pollock) but are more widely distributed throughout the tissues of piscivorous fish (whiting, cod). These observations are discussed in relation to the apparent importance of euphausiids as intermediate hosts of A. simplex.