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.     

Hatching rates of resting eggs of 'Cladocera' (Crustacea; Branchiopoda) at a tropical bay, Brazil.

The aim of this study was to determine the development time of embryos and to estimate the hatching rates of resting eggs of cladocerans found in the sediment of Guanabara Bay, Rio de Janeiro, Brazil, under experimental conditions. Eggs were sorted by species (Penilia avirostris--Sididae; Pleopis polyphemoides and Pseudevadne tergestina--Podonidae) and incubated at a temperature of 25 degrees C, salinity 35 and photoperiod 12 hours light/ 12 hours dark. Hatching rates were about 38% for Pseudevadne tergestina and 28% for Pleopis polyphemoides. Embryos of resting eggs of Penilia avirostris developed comparatively slowly (hatching after 86 days of incubation), with a hatching rate of only 5%. It was observed that development and hatching of resting eggs of marine cladocerans suggest that pulses of recruitment may exist, thus contributing to the rapid appearance and maintenance of planktonic populations of these crustaceans in Guanabara Bay.     

Sex-biased egg cannibalism in spawning walleye pollock: the role of reproductive behavior

We studied inadvertent egg cannibalism in spawning stocks of walleye pollock, Theragra chalcogramma, in the Gulf of Alaska and the Bering Sea between 1986 and 1996. Male pollock had on average 3 times more eggs in their stomachs than females. For both sexes adult fish of average body length had more eggs in their stomach than did smaller or larger fish. When maturity of fish was taken into account, actively spawning males had the highest numbers of eggs in their stomachs. We found weak evidence for a diel variation; the number of eggs per stomach was high in both sexes during the day and lower during the night. We suggest two possible explanations for this phenomenon. Sex-biased egg cannibalism may reflect the differential time spent in layers of high egg densities. Hydroacoustic and trawl catch data from both areas suggest that males aggregate deeper than females. Spawning takes place in the deeper layers of the fish aggregation, so males spend more time in high egg densities. Alternatively, males may be more active than females and increased gill ventilation and/or drinking rates may be responsible for the differences in egg cannibalism.     

Ontogenetic shift in geotaxis for walleye pollock,Theragra chalcogramma free embryos and larvae: potential role in controlling vertical distribution

Synopsis The behavioral capability of walleye pollock,Theragra chalcogramma free embryos and larvae to control vertical distribution was assessed by examining buoyancy during resting and swimming orientation and activity as they developed in complete darkness from hatching to first feeding readiness (1 to 7 d post hatching at 6° C). Free embryos exhibited positive geotaxis 1 d post hatching, actively swimming through a density gradient to remain in the lower water column. Activity increased with free embryo development and by 7 d post hatching, feeding-ready larvae reversed their vertical orientation, now exhibiting negative geotaxis as they migrated to the upper water column. The results indicate that even at the earliest developmental stages, walleye pollock possess the capability to control vertical distribution. Laboratory results are compared with patterns of vertical distribution observed in the sea.     

Assessment of the consumption of walleye pollock eggs by nekton and jellyfish in the northern Sea of Okhotsk in the spring

In the northern Sea of Okhotsk, nekton and jellyfish consumed as many as 831 × 10⁹ walleye pollock eggs per day in 2011. The nekton exerted the highest pressure, viz., 98.3% of the overall predation on pollock egg by aquatic animals. Of the entire quantity of consumed eggs, 55.9% were eaten by herring, 35.9% by walleye pollock, 6.5% by Sakhalin sole, and 1.7% by jellyfish. Among jellyfish, scyphomedusae Cyanea capillata and Chrysaora melonaster, as well as the hydromedusa Tima sachalinensis consumed the largest quantities of eggs. The total consumption of pollock egg by aquatic animals in 2011 was estimated at 42.4 × 10¹², or 11.4% of the entire quantity of eggs that were spawned by walleye pollock in the waters of the northern part of the sea. The total amount of pollock eggs that were eaten by herring and pollock together for 51 days in 2011 amounted to 38.9 × 10¹², which was 5.7 times as much as that in 2002. Thus, a significant growth of predation on pollock eggs by their main consumers, viz., herring and walleye pollock, was observed in 2011. This was caused by an increase in the populations of both species during the recent years and also by a higher concentration of pollock eggs.     

Circadian clock controlling egg hatching in the cricket (Gryllus bimaculatus)

Adult crickets (Gryllus bimaculatus) were maintained under a 12-h light:12-h dark cycle (LD 12:12). After oviposition, their eggs were incubated under different lighting regimens at 23 degrees C, and temporal profiles of egg hatching were examined. When the eggs were incubated in LD 12:12 or in DL 12:12 with a phase difference of 12h from LD 12:12, throughout embryogenesis, 88% to 97% of hatching occurred within 3 h of the dark-light transition on days 17 and 18 of embryogenesis; the phases of the egg-hatching rhythms in the LD 12:12 and DL 12:12 groups differed by about 12 h. In eggs incubated in constant darkness (DD) throughout embryogenesis, a circadian (about 24 h) rhythm of hatching was found, and the phase of the rhythm was similar to that seen in eggs incubated in LD 12:12, but not DL 12:12, throughout embryogenesis. When eggs that had been incubated in DD after oviposition were transferred to DL 12:12 in the middle or later stages of embryogenesis and were returned to DD after three cycles of DL 12:12, the rhythm of hatching synchronized (entrained) to DL 12:12. However, when eggs in the earlier stages of embryogenesis were transferred from DD to DL 12:12 and returned to DD after three cycles, 52% to 94% of hatching did not entrain to DL 12:12. To determine whether photoperiodic conditions to which the parents had been exposed influenced the timing of egg hatching, adult crickets were maintained in DL 12:12, and their eggs were incubated in LD 12:12, DL 12:12, or DD throughout embryogenesis. The egg-hatching rhythm was also found in the eggs incubated under these three lighting regimens. In DD, the phase of the rhythm was similar to that seen in eggs incubated in DL 12:12, not LD 12:12, throughout embryogenesis. The results indicate that in the cricket, the timing of egg hatching is under circadian control and that the circadian rhythm of hatching entrains to 24-h light:dark cycles, but only if the light:dark cycles are imposed midway through embryogenesis. Therefore, by midembryogenesis, a circadian clock has been formed in the cricket, and this is entrainable to light:dark cycles. In addition, the photoperiodic conditions to which the parents (probably the mothers) have been exposed influence the timing of hatching, suggesting that maternal factors may regulate the timing of egg hatching.     

Effects of oxygen,carbon dioxide,and nitrogen on hatching behavior and hatching time in the katydid,Eobiana engelhardti subtropica Bey-Bienko (Orthoptera: Tettigoniidae)

The trigger for the hatching behavior and determination of hatching time of the katydids, Eobiana engelhardti subtropica (Orthoptera: Tettigoniidae) have been shown to be influenced by light–dark signals or temperature. In this study, I investigated the effects of oxygen, carbon dioxide, and nitrogen on the hatching behavior and hatching time of the katydid. Eggs rarely hatched under a constant temperature of 25°C and hatched sporadically at a constant temperature of 15°C under continuous light in the air. However, when eggs were exposed to 100% oxygen or a mixture of oxygen and nitrogen (2:1 or 1:1), hatching occurred within a few seconds. Hatching behavior was directly triggered by high concentrations of oxygen. It was inhibited by exposure to 100% carbon dioxide, 100% nitrogen, or a mixture of oxygen and nitrogen (1:2). The hatching time, determined by the temperature fall (transfer from 25°C to 15°C), was delayed by these gases, and was reset by the transfer back of eggs to the air. This suggests the existence of a time-measuring mechanism that is triggered by the transfer of eggs to the air. These results, indicating that hatching behavior was directly triggered by high concentrations of oxygen and that hatching time was set by the transfer from carbon dioxide or nitrogen to the air, are new findings to the best of my knowledge.     

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.     

Development and application of a bioenergetics model for juvenile walleye Pollock

A bioenergetics model was parameterized for age-0 walleye pollock, Theragra chalcogramma , based on a synthesis of literature data. The sensitivity of the new parameters was tested by individual parameter perturbation (IPP) analysis. The model was applied to estimate individual and total cohort food consumption of age-0 pollock in two areas of high pollock density in the Gulf of Alaska during the summer of 1990. Total cohort consumption was compared with zooplankton biomass and production estimates for the same areas and times of the year. The model was also used to examine the bioenergetic implications of age-0 pollock diel vertical migration through a thermal gradient. During a 1-month bioenergetics simulation, individual daily consumption decreased from 16·0 to 6·0% of wet body weight. Daily ration estimates corresponded well with independent field estimates of daily ration for the same areas and time of the year. Comparison of total cohort consumption with prey availability (production and biomass) indicated minimal potential for food limitation. Bioenergetic optimization of growth can be a potential benefit of diel vertical migration to age-0 pollock, however more information on prey density and distribution is needed to test this hypothesis thoroughly.     

Hatching rhythms in three species of Diclidophora (Monogenea) with observations on host behaviour.

Eggs of three species of Diclidophora were incubated in alternating 12 h periods of light and darkness at 13 degrees C. Eggs of D. merlangi collected at Arbroath hatched during the illumination period with most larvae being recovered in the first 4-6 h; some evidence of a seasonal difference in hatching of these eggs was found. Eggs of D. merlangi collected at Plymouth hatched with a peak of larval recovery in the 2 h period before the light came on. Eggs of D. luscae hatched over 'dusk' while those of D. denticulata hatched after the light was switched off. Neither mechanical disturbance nor the proximity of host tissue caused hatching in D. merlangi or D. luscae. Observations on the behaviour of the host fishes suggest that the hatching rhythms are adapted to specific host behaviour patterns.     

Changes in mass of eggs of softshell turtless (Trionyx spiniferus) incubated under hydric conditions simulating those of natural nests

Rigid-shelled eggs of softshell turtles ( Trionyx spiniferus ) incubated at 29C under hydric conditions simulating those at the interior of natural nests (i.e. eggs not contacting the substrate) declined in mass during incubation owing to the continuous escape of water vapour to air trapped inside the chambers, with eggs incubated in relatively dry chambers declining more in mass than eggs held in relatively wet chambers. Eggs held under hydric conditions approximating those at the periphery of natural nests (i.e. eggs partly buried in the substrate) also declined in mass during incubation, but at lower rates than characterized eggs not contacting the substrate, indicating that absorption of liquid water from the substrate compensated for part of the water vapour escaping to the air inside the chambers. There was no relationship between hatching success and either the position of eggs in the chambers or the wetness of substrates, nor was there any variation among experimental groups in the size of hatchlings. Thus, development of embryos of Trionyx spiniferus seems not to be closely coupled with the hydric environment of the nest, a situation contrasting with that of embryos of those species of turtles producing pliable-shelled eggs.     

Field and laboratory studies on the timing of oviposition and hatching of the western black-legged tick, Ixodes pacificus (Acari: Ixodidae)

The timing of oviposition and hatching of Ixodes pacificus was investigated in the field and at constant temperatures in the laboratory. Replete females held at temperatures between 9 and 29°C began depositing eggs a mean of 9–70 days after drop off. Egg masses held between 12 and 25°C commenced hatching 25–178 days after the onset of oviposition. Eggs held at 9 or 29°C did not hatch. The lower temperature thresholds for development (LTD) for oviposition and hatching were 6.5 and 9°C, respectively. The number of degree days required for oviposition and hatching was 173 and 588, respectively. Replete females placed in the field on 2 December through to 8 March deposited eggs from 2 February through to 24 April; the eggs commenced hatching between 2 July and 21 August. Unfed larvae from two of 20 egg masses survived through the winter and fed readily when exposed to deer mice (Peromyscus maniculatus) on 22 April. Replete larvae were returned to the field and moulted between 9 and 21 August. Larvae exposed to deer mice in August, 4 weeks after hatching, also fed readily. Although further studies are needed to clarify the timing of nymphal development, the present study suggests that I. pacificus requires more than 1 year to complete its life cycle.     

Spatial Match-Mismatch between Juvenile Fish and Prey Provides a Mechanism for Recruitment Variability across Contrasting Climate Conditions in the Eastern Bering Sea

Understanding mechanisms behind variability in early life survival of marine fishes through modeling efforts can improve predictive capabilities for recruitment success under changing climate conditions. Walleye pollock (Theragra chalcogramma) support the largest single-species commercial fishery in the United States and represent an ecologically important component of the Bering Sea ecosystem. Variability in walleye pollock growth and survival is structured in part by climate-driven bottom-up control of zooplankton composition. We used two modeling approaches, informed by observations, to understand the roles of prey quality, prey composition, and water temperature on juvenile walleye pollock growth: (1) a bioenergetics model that included local predator and prey energy densities, and (2) an individual-based model that included a mechanistic feeding component dependent on larval development and behavior, local prey densities and size, and physical oceanographic conditions. Prey composition in late-summer shifted from predominantly smaller copepod species in the warmer 2005 season to larger species in the cooler 2010 season, reflecting differences in zooplankton composition between years. In 2010, the main prey of juvenile walleye pollock were more abundant, had greater biomass, and higher mean energy density, resulting in better growth conditions. Moreover, spatial patterns in prey composition and water temperature lead to areas of enhanced growth, or growth ‘hot spots’, for juvenile walleye pollock and survival may be enhanced when fish overlap with these areas. This study provides evidence that a spatial mismatch between juvenile walleye pollock and growth ‘hot spots’ in 2005 contributed to poor recruitment while a higher degree of overlap in 2010 resulted in improved recruitment. Our results indicate that climate-driven changes in prey quality and composition can impact growth of juvenile walleye pollock, potentially severely affecting recruitment variability.     

Evidence for embryo–embryo interactions in controlling hatching time and synchronization in the brown marmorated stink bug,Halyomorpha halys (Hemiptera: Pentatomidae)

The mechanisms controlling egg diapause and circadian rhythms of hatching activity have been extensively studied in insects. However, relatively little attention has been paid to the mechanisms controlling synchronized hatching from an egg mass. In this study, we examined the possible involvement of embryo–embryo interaction in controlling hatching time in Halyomorpha halys (Stål). Eggs tended to hatch earlier as the egg mass size increased. Egg separation and clumping of separated eggs at various times showed that hatching synchrony was largely determined shortly before hatching. However, whether eggs were kept in a mass or separated until several hours before hatching also influenced the hatching time, indicating the presence of embryo–embryo interactions. Eggs derived from different masses and kept in physical contact with one another hatched synchronously if their ages were within approximately 8 h. In this case, both younger and older eggs advanced only in hatching time, in contrast to a case of locusts reported by others. Eggs separated by more than 7 mm hatched as synchronously as those kept in a mass when glued to the same substrate, suggesting an important role of the egg substrate in transmitting the vibrational hatching signals to neighboring sibling eggs to synchronize hatching.     

Effects of light on pigment development in the echiuran worm, Bonellia viridis

In rhe echiuran Bonellia viridis the green pigment first develops in the gastrula, before the formation of the trochal rings, and the trochophores acquire a deep green pigmentation before hatching. Eggs that were cultured under continuous illumination (2000–4000 lx) did not develop any green pigment, and although the trochophores hatched normally, they lacked pigment entirely. The unpigmented trochophores that were kept under continuous illumination developed into unpigmented males and females. The far-blue (380–400 nm) and red (620–640 nm) regions were more effective in preventing pigment development than the blue, green and yellow regions of the spectrum. Illumination of the eggs before the gastrula stage did not affect the development of pigment when these were subsequently returned to the dark. When unpigmented trochophores, that had been cultured in the light, were placed in the dark they acquired a slight pigmentation, but this was much less intense than that of trochophores that had been cultured in the dark throughout. Pigmented trochophores were more vulnerable to high light intensities than unpigmented ones.     

Diel periodicity of emergence of adult citrus rust mites in central Florida

Emergence of adult citrus rust mites, Phyllocoptruta oleivora (Ashmead) from the imagochrysafis occurred with a diel periodicity that peaked between mid morning and early afternoon. The pattern of emergence did not differ for mites reared from eggs held under different photoperiod regimes. The transition of nymphs to the imagochrysalis stage also showed a diel periodicity, occurring in the evening. The average duration of the imagochrysalis stage at 28 °C was 14–15 h.Wetting by rain or dew was simulated by immersing fruit in water each night, but the transition to the imagochrysalis stage or the subsequent emergence pattern of adults was not affected. Adult mites reared from eggs laid during two, consecutive 12-h periods emerged at the same time of day, but the peaks of emergence were separated by 24 h. Eclosion of larvae from eggs laid during two, consecutive 12-h periods did not show a diel periodicity. Eggs continuously immersed in water during their development did not hatch, whereas there was a small, but significant, reduction in the viability of eggs immersed in water each night. These results are discussed in relation to external cues that may entrain the diel patterns of development and adult emergence, and to the adaptive responses of citrus rust mite to cyclic fluctuations of key environmental variables in central Florida. This revised version was published online in August 2006 with corrections to the Cover Date.     

Abiotic factors influencing embryonic development, egg hatching, and larval orientation in the reindeer warble fly, Hypoderma tarandi

Wild-caught, tethered females of the reindeer warble fly, Hypoderma tarandi (L.) (= Oedemagena tarandi (L.)), (Diptera, Oestridae) were stimulated to oviposit on hairs of a reindeer hide. Newly laid eggs incubated at constant temperatures and relative humidities hatched within 3 days to 2 weeks, depending on the experimental conditions. Over a range of 7-40 degrees C, hatching only occurred between 20 and 37 degrees C. Eggs held at 100% relative humidity had lower hatchability and longer time to hatch relative to eggs held at 77% relative humidity. The average number of degree-days for hatching was 50.35. Between 20 and 33 degrees C there was a temperature-dependent linear trend in developmental rate, and the proportion of eggs hatching was highest, and least variable, at the mid-temperature ranges. The temperature range found in the natural host micro-habitat where H. tarandi commonly affix their eggs (close to the skin at the base of a host hair) was consistent with the experimental temperature treatments that produced the highest hatching rate. Newly emerged larvae displayed positive thermotaxis, while showing no phototaxic or geotaxic behaviour. Results indicate that constraints of the host environment, coupled with temperature-dependent hatching success, may impose a selective pressure on oviposition behaviour.     

Diel activity pattern and predation rate of the generalist predator Dicyphus hesperus

Abstract This study examined the diel activity pattern and the effect of diel activity pattern on predation rate and prey finding of Dicyphus hesperus Knight (Heteroptera: Miridae). To determine the diel activity pattern of D. hesperus, starved females were placed on tomato leaflets Lycopersicon esculentum Mill. (Solanaceae) under zero, low, or high light intensities at 02:00, 08:00, and 14:00 h, respectively, and the amount of time spent walking or resting during a 30‐min interval was recorded. Predation rates of D. hesperus females on Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) egg patches were determined under either a L16:D8 (long day) or L8:D16 (short day) diel period. Egg patches were removed from D. hesperus females after either 8 or 16 h of dark or 16 or 8 h of light, and the number of eggs consumed was counted. Dicyphus hesperus females spent more time searching for prey at night than during the day. Females ate eggs at a higher rate during the night than during the day. Overall, D. hesperus females had higher predation rates when reared under a long day diel cycle compared with females reared under a short day diel cycle. More females reared at the L16:D8 diel cycle found the egg patch during the night than during the day. There was no difference in egg patch finding between night and day for females reared at L8:D16. Overall, L16:D8 reared females found more egg patches than females reared at L8:D16. Therefore, D. hesperus females are more active and find and consume prey at a higher rate at night than day.     

Effects of simulated trawling on sablefish and walleye pollock: the role of light intensity, net velocity and towing duration

Laboratory apparatus which simulated capture of fish in the cod-end of a towed trawl was used to induce post-capture stress as measured by alterations in behavioural, physiological and mortality indices in juvenile walleye pollock Theragra chalcogramma and juvenile and adult sablefish Anoplopoma fimbria. Differences in resistance to net entrainment varied between species with the severity of stress and the potential for recovery depending on light intensity, net velocity and towing duration. At a light intensity which simulated daylight at depth in clear ocean water (0.5 μmol photons m^?2 s^?1), walleye pollock juveniles were able to maintain swimming in nets towed at 0.65 m s^?1 for 3h with no discernible effects on behaviour or mortality. However, when net velocity was increased to >0.75m s^?1 or light intensity was decreased to <0.002 μmol photons m^?2 s^?1, fish became entrained in the meshes of the net and exhibited significant alterations in feeding behaviour, predator evasion and increases in plasma cortisol concentrations. Marked increases in stress-induced mortality also occurred, in some cases after a delay of 6 days and eventually reaching 100%. In comparison with walleye pollock, sablefish juveniles became entrained in the meshes of the net at higher velocities (>0.92m s^?1) or lower light intensities (<0.0004 μmol photons m^?2 s^?1) and were much more resistant to post-capture stress. Towing of net-entrained fish for 15 min caused no detectable changes in feeding and cortisol and for 2 h, no changes in feeding although mortality increased from 0% for 15-min tows to 19% for 2-h tows. Towing for 4 h caused significant alterations in feeding and cortisol with feeding recovering to control levels by 6 days and cortisol by 3 days; mortality was 25%. When adult sablefish were towed for 4 h followed by 15-min exposure to air, feeding was inhibited 6 days after towing, but recovered within 30 days with no mortality observed after 30 days. The results demonstrate the value of using laboratory-based behavioural and biochemical indices to identify factors that may potentially affect post-capture survival among different species of fish.     

Accessibility of sulfhydryl groups to 5,5'-dithiobis-2-nitrobenzoic acid and acid-base properties of bovine and walleye pollock rhodopsin preparations]

Both the number of exposed SH-groups and the rate of reaction with 5,5'dithiobis-2-nitrobenzoic acid (DTNB) in walleye pollock and bovine rhodopsin depend on a degree of native structure of the preparation to be investigated. The preparations studied can be arranged in the order of increase of these parameters as follows: ROS less than rhodopsin extracted by digitonin less than triton X-100 less than cetyltrimethylammonium bromide (CTAB) less than sodium dodecylsulphate (SDS). After illumination of ROS and digitonin, triton X-100 and CTAB-solubilized rhodopsin, and increase was observed in the number of modified SH-groups. Dark and bleached samples of walleye pollock rhodopsin exhibited a faster rate reaction and a more number of modified SH-groups as compared to bovine preparation. The differences between bovine and walleye pollock preparation disappeared after complete opsin unfolding as a result ROS solubilization in SDS. Six SH-groups per molecule of rhodopsin were modified in both preparation under these conditions. No differences in the number of cysteine residues (10--11), disulfide groups (2), acid (35--40) and base (25--30) titratable groups per rhodopsin molecule were found between bovine and walleye pollock ROS membranes. The isoelectric point of both rhodopsin preparations was within the pH range 5.2--5.6. After proteolysis of ROS with papain, a fragment with molecular weight 24500 +/- 1000 was detected, which contained the same number of SH-groups and cysteine residues as in the case of intact rhodopsin. The results obtained suggest that, in spite of a similar primary structure, the walleye pollock visual pigment has more "loose" and "fluid" space packing in the ROS membrane than the bovine pigment.