Ontogenetic development of digestive enzyme activities in larval walleye pollock,Theragra chalcogramma

Activities of digestive enzymes trypsin, amylase and lipase in laboratory-reared walleye pollock, Theragra chalcogramma, were measured from hatching to Day 39 (just before notochord flexion) in 1993. All measurements were conducted individually or semi-individually (groups of two larvae of the same standard length). Close relationships between digestive enzyme activities and morphological development of digestive organs were observed. Activities of trypsin and lipase were low during the transition period from endogenous to exogenous energy. Amylase activity was constant with large variance during the same period. Specific enzyme activities of trypsin and amylase indicated high values with large variance during the early period. All three enzyme activities increased with age afterthe transition period, and the specific enzyme activities became constant. The existence of two types of lipase was suggested. One lipase showed a peak of specific activity at Day 4 and might be related to yolk-sac absorption. The activity of the other lipase increased with age after Day 14 and might be related to digestion of prey lipid. Our results suggest that digestive enzymes included in food organisms supplement larval pollock digestive enzymes.     

Evaluating growth of larval walleye pollock, Theragra chalcogramma, using cell cycle analysis

Cell cycle analysis of muscle cell division rates offers a new and efficient technique to analyze growth of larval fish. Using this approach, growth of larval walleye pollock was estimated by determining cell proliferation rates, reasoning that growth during early life stages is probably attributed to increases in cell number rather than to increases in cell size. Characteristic patterns of brain and muscle cell division rates were produced in larval walleye pollock by manipulating their diet in the laboratory. The fraction of dividing muscle cells and, to a lesser extent, the fraction of dividing brain cells were direct indicators of fast and slow growth. A model was produced to estimate average growth rate from the fraction of dividing muscle cells. We developed a simple method for preparing and storing the muscle tissue that ensures nucleic acid stability for subsequent analyses and permits sampling in the field. We envision that the cell cycle methodology will have on-site applications, presenting an opportunity to attain real-time estimates of larval fish growth at sea. Determining the proportion of first-feeding larvae with a high fraction of dividing muscle cells may yield a means for predicting the proportion of fast-growing fish, i.e., the potential survivors.     

Otolith analysis of juvenile walleye pollock Theragra chalcogramma from the western Gulf of Alaska

Growth rate and hatch date distributions were estimated for juvenile walleye pollock Theragra chalcogramma (Pallas, 1814) collected in autumn 1987 from the western Gulf of Alaska. Mean juvenile growth rates varied geographically by as much as 45%. A trend for slower growth around Unimak Pass and the Shumagin Islands and faster growth upstream in the Alaska Coastal Current towards Kodiak Island was noted; the fastest growth did not occur in the main habitat region. Juvenile hatch date distributions were compared to identify regional differences that might reflect stock structure. Juvenile hatch dates were compared with hatch dates of the same cohort sampled as larvae to test for selective mortality. Regional differences in hatch dates, along with other information, indicated several minor spawning populations located around Kodiak Island and near Unimak Pass. For the main aggregation of pollock in the Shumagin Island region, hatch date distributions were not significantly different among the early larval cohort sampled in late May, the late larvae sampled in mid-June to early July, and the juveniles sampled in autumn. Neither growth-rate nor size-dependent mortality of pollock between the larval and juvenile stages appears to be a dominant factor in determining survival patterns. An alternative test was attempted, by which lengths-at-age during larval life were back-calculated from juvenile otoliths and compared with lengths-at-age of the population sampled as larvae in May and June. Pollock surviving as juveniles in autumn were not larger as larvae than the general larval population. This result is an example of the observation that back-calculated lengths are almost always smaller than the lengths of fish sampled at age (Lee's phenomenon). Problems in determining survival patterns based on otolith back-calculations and comparison of hatch date distributions are discussed.     

Underlying causes of habitat-associated differences in size of age-0 walleye pollock (Theragra chalcogramma) in the Gulf of Alaska

Age-0 walleye pollock (Theragra chalcogramma) caught in September in the Gulf of Alaska display habitat-associated differences in standard length (SL). Age-0 fish collected in the region around Sutwik Island and 375?km farther downstream near the Shumagin Islands most likely originate from the Shelikof Strait spawning aggregation. However, age-0 fish resulting from the same spawning aggregation differ in mean size up to 20?mm between areas by September. We examined the otoliths of the larval and age-0 stages of walleye pollock from these two areas in 2000 and 2001 to determine whether growth rate, hatch date, and/or temperature influenced fish size. Circulation models were used to determine whether transport of larvae from an upstream spawning group into the study areas could have occurred. Mean in situ temperature during sampling periods was not defined as a significant factor in altering growth rates. Overlapping hatch date distributions of the larval and age-0 fish in the Shumagin Island area confirmed that the fish were from the Shelikof Strait spawning group. Comparison of hatch date distributions in the upstream Sutwik Island area revealed larger/older larvae from an upstream spawning group mixed with larvae from the Shelikof Strait spawning group. Our results suggest that the offset of 20?mm SL between the groups of age-0 pollock was the result of a combination of enhanced survivorship of early-hatched larvae in the Sutwik area and the introduction and retention of the progeny of another spawning group originating upstream of Shelikof Strait.     

Predation on walleye pollock (Theragra chalcogramma) eggs in the western Gulf of Alaska: the roles of vertebrate and invertebrate predators

Estimates were made of the predation rate upon eggs of walleye pollock (Theragra chalcogramma) in Shelikof Strait in the western Gulf of Alaska by midwater and near-bottom fish and invertebrate predators during April 1990. Adult and juvenile walleye pollock were the dominant (99% of total abundance) planktivores collected in midwater samples. Based on visual inspection of stomach contents, a high percentage of the sampled fish were found to have consumed pollock eggs. Daily egg consumption by older age groups of walleye pollock was estimated to be <1% of the eggs available at all sampling locations. The only other fishes found to consume pollock eggs were flatfishes collected in bottom trawls but their abundances and egg consumption were very low. Gammarid and hyperiid amphipods were important invertebrate predators on eggs in the water column, as determined by immunoassays using antibodies developed specifically to ascertain the presence of pollock egg-yolk protein. Decapod shrimp showed a high proportion of positive assays in near-bottom collections. Invertebrate predators may have consumed up to 4% of the total number of eggs available in the water column, but <1% of the total near the bottom on a daily basis. Although we were not able to account for the entire daily egg mortality estimated for this stock, our method of using a combination of techniques is promising in terms of future attempts at estimating total predation mortality.     

Geographical differences in early growth of walleye pollock Theragra chalcogramma,estimated by back-calculation of otolith daily growth increments

The relationship between otolith length and body length (total length) was examined in walleye pollock Theragra chalcogramma (Pallas), collected from Pacific Ocean waters adjacent to Hokkaido, Japan, and in larvae reared from fertilized eggs. A linear relationship was found between log-transformed body length and otolith length data with two inflection points at ca. 11 mm and 100 mm total length. This relationship was found to be applicable also to samples from the Japan Sea, Bering Sea and Okhotsk Sea. The early growth pattern estimated by back-calculation of otolith increments of 1-yr fish from Funka (Uchiura) Bay (Hokkaido) accorded with that obtained from size-at-age data of 0 yr fish collected in the preceding year. Differences were found in comparisons of the back-calculated early growth pattern between samples from waters adjacent to Hokkaido, the Pacific Ocean, the Japan Sea and the Okhotsk Sea. The growth curve of the Okhotsk samples was markedly different from others, showing rapid initial growth up to about 100 d after hatching and attaining a small body size in the first year.     

Foraging impact on zooplankton by age-0 walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) around a front in the southeast Bering Sea

The waters around the Pribilof Islands in the southeast Bering Sea are a center of abundance for age-0 walleye pollock (Theragra chalcogramma). Each spring and summer a tidal front is formed around the islands separating a well-mixed inshore habitat from a stratified offshore habitat. The objective of this study was to assess the foraging impact on zooplankton by age-0 pollock in the vicinity of this frontal structure. A bioenergetic model was used to estimate age-0 pollock food consumption from field estimates of water temperature, age-0 pollock density, diet and growth. Sampling of field variables took place over three hydrographic habitats along an inshore–offshore transect located north of the islands. The bioenergetics analysis was applied for a 2-week period during the late summer of four consecutive years, 1994–1997. Model results of age-0 pollock food consumption indicated variable levels of food depletion, changing with prey type, year and habitat. The foraging impact of age-0 pollock on copepods and euphausiids (most common prey) ranged from about 3% to 77% of the biomass available at the start of the simulation. Copepod depletion was typically greater than euphausiid depletion. Consequently, juvenile pollock <60 mm in standard length were more likely to experience food limitation due to the greater proportion of copepods in their diet. We present evidence of severe foraging impact during 1996, when one of the primary prey items of juvenile pollock (i.e. large copepods) was scarcely represented both in their diet and in the water column. In all years, most instances of prey depletion were found at the inshore and front habitats; age-0 pollock densities were too low relative to their prey to severely impact the offshore zooplankton populations. We discuss these results with respect to modeling assumptions and in the context of previously acquired knowledge of fish behavior around frontal regions.Communicated by T. Ikeda, Hakodate     

Effect of temperature on the escape responses of larval herring,Clupea harengus

Herring (Clupea harengus L.) larvae from spring and autumn spawning stocks were reared at different constant temperatures from 5° to 17 °C. At equivalent developmental stages, the spring larvae were longer than the autumn larvae and the larvae reared at low temperatures were longer than those reared at high temperatures. At hatching and at the end of the yolk-sac stage, the larvae were induced, by a probe, to make C-start escape responses, which were recorded and analysed using a high-speed video recording at 400 frames s^-1. The response was rapid and of short duration. The tailbeat frequency and swimming speed were measured during the burst of swimming following the C-start at different test temperatures and in larvae with different temperature histories. The tail-beat frequency was strongly temperature-dependent, rising from 19 Hz at 5 °C to 37 Hz at 17 °C with no effect of temperature history, season or developmental stage. The burst-swimming speed ranged at hatching from 75 to 90 mm s^-1 at 5 °C to 110 to 160 mm s^-1 at 17 °C and at yolk resorption from 90–115 mm s^-1 at 5 °C to 175–190 mm s^-1 at 17 °C. The longer, spring-spawned larvae swam faster than the shorter autumn-spawned larvae. When the swimming speeds were expressed as body lengths (L) s^-1, these differences disappeared. Larvae swam from 7–9 L s^-1 at 5 °C to 15–20 L s^-1 at 17 °C at hatching, and from 8–9 L s^-1 at 5 °C to 15–17 L s^-1 at 17 °C at yolk resorption. There was, however, a significantly faster specific swimming speed by the larvae reared at 12 °C in spring 1991.Honorary Research Fellow of the Scottish Association for Marine ScienceUnfortunately, Karen Fretwell was drowned in an accident on 9 January 1993     

Switching to Plan B: changes in the escape tactics of two grasshopper species (Acrididae: Orthoptera) in response to repeated predatory approaches

Most studies examining escape behaviour have considered single approaches and single fleeing responses; few have considered how organisms’ response is influenced by persistent pursuit. We explored fleeing behaviour of two grasshopper species to test whether they modified escape behaviour when approached repeatedly. Schistocerca alutacea did not increase flight initiation distance (FID) upon repeated approach but fled farther. Psinidia fenestralis increased its FID on the second approach but decreased its flight distance over successive escapes. Both species showed a bimodal pattern of flight direction, either flying directly away or flying perpendicular to the direction of the observer’s approach. Neither species showed a significant pattern of flight direction or change in flight direction with successive escapes. Most (88 %) P. fenestralis initially landed on sand, but after repeated approaches an increasing proportion landed in grass and hid. Both species therefore changed escape behaviour with persistent pursuit but used different tactics, suiting their flight ability or camouflage, and optimised habitat use. Three grasshopper species have now been examined for responses to repeated approach by predators and all show different tactics supporting escape decision theory. Our results emphasise the variety of escape responses across species and how the dynamic nature of escape responses vary according to an animal’s situation. Rather than single optimum escape options, each grasshopper species shows a range of responses, which vary with risk from persistent predators. Although grasshoppers provide an excellent model, it would be profitable to examine responses of a range of species according to levels of predation risk.     

Calanoid copepod escape behavior in response to a visual predator

Calanoid copepods typically exhibit escape reactions to hydrodynamic stimuli such as those generated by the approach of a predator. During the summers of 2000, 2001 and 2004, two small calanoid species, Temora turbinata Dana, 1849 and Paracalanus parvus Claus, 1863 were exposed to a visual predatory fish, the blenny Acanthemblemaria spinosa Metzelaar, 1919, and their predator–prey interactions were recorded using both high-speed and standard videographic techniques. Copepod escape reaction components, including swimming pattern, reactive distance, turning rate, and jump kinetics, were quantified from individual predation events using motion analysis techniques. Among the observed escape reaction components, differences were noted between the species’ swimming patterns prior to attack and their response latencies. Temora turbinata was a continuous cruiser and P. parvus exhibited a hop-and-sink swimming pattern. During periods of sinking, P. parvus stopped beating its appendages, which presumably reduced any self-generated hydrodynamic signals and increased perceptual abilities to detect an approaching predator. Response latency was determined for each copepod species using a hydrodynamic stimulus produced by a 1 ms acoustic signal. Response latencies of T. turbinata were significantly longer than those of P. parvus. Despite some apparent perceptual advantages of P. parvus, the blenny successfully captured both species by modifying its attack behavior for the targeted prey.     

Development of the pH in the intestinal tract of larval turbot

The pH in the gut of turbot larvae and juveniles of turbot was studied from day 11 until the completion of metamorphosis. Dietary effects on the gut pH were estimated when larvae were offered live feed, a microdiet, only microalgae or no feed. The pH in the gut was weakly alkaline until day 24 after hatching with no differences between the foregut, midgut and hindgut. The foregut contents started to turn acidic from day 28 after hatching when the larvae were already weaned successfully, which indicates that an acidic pH is not necessary for the digestion and utilisation of formulated feed. During the following 20 days the pH in the foregut/stomach decreased further to a minimum of pH 3.5, while the pH in the midgut and the hindgut increased slightly to a maximum of pH 9.0. Larvae receiving live feed, microdiet or microalgae had a similar pH in the midgut on day 11, while starved larvae exhibited a lower gut pH. This suggests bicarbonate secretion from the larval pancreas stimulated by ingested microalgae or feed particles.     

The larval settling response of the lined chiton Tonicella lineata

The lined chiton Tonicella lineata (Wood, 1815) is found on enerusting coralline algae in the lower rocky intertidal zone of Oregon and San Juan Island, Washington, USA. An encrusting coralline alga is the major food item of this chiton. Experiments were performed to test the settling response of T. lineata larvae to various algae and other substrata. In these experiments, the larvae would settle only on pieces of encrusting coralline algae and piecesof ceramic roofing tile soaked in a coralline algal extract. The settling stimulus is probably chemical in nature, and is inactivated by boiling. In laboratory cultures, normal development stops at the trochophore stage (110 to 160 h, depending on the temperature). Metamorphosis and further development will take place only after stttlement on encrusting coralline algae.     

Dominance of larval forms in euphausiid (Crustacea:Eucarida) ontogenesis

Variability and dominance of Euphausia superba Dana larval forms (Furcilia-I and Furcilia-II stages) in the plankton of the Scotia and Weddell Seas have been investigated. New forms (1, 2, 3 and 14) have been recorded. The dominant forms of Furcilia-I stage are proposed to represent the initial dominants. In accordance with the initial dominance, five types of larval form combinations have been established. During the initial dominance change — from 5 to 2, the larval form number increases. This phenomenon is due to parallel development of euphauiid larvae; it results from a developmental pathway change in the main larval mass. Such deviations are reflected in an initial dominance change. In each pathway, the larvae pass through an unequal number of moults before reaching Form 5 (in accordance with the proposed moult scheme of Furcilia-I stage larvae). In the case of initial dominance of less-developed forms, the presence of different parallel pathways increases. As a result, the total number of larval forms present in the plankton changes, causing increased variability of the total number of larval forms. Each type of larval form combination is the result of summation of all larval forms, which derive from different developmental pathways of the larvae in different localities. The most usual developmental pathway for Euphausia superba is through Form 5, which may, thus, be regarded as the main developmental pathway for this species. This pathway probably corresponds to optimal environment conditions for larval growth and morphogenesis with maximal synchronisation of both processes. Consequently, it may be suggested that changes in developmental pathways of larvae are connected with suboptimal environmental conditions.     

Quantitative genetic analysis of growth in larval scallops (Placopecten magellanicus)

We estimated the broad-sense heritability of larval size in 20 full-sib families of the giant scallop, Placopecten magellanicus (Gmelin, 1791) grown in laboratory culture in August and September 1991. The goal was to compare scallops with other bivalves which have been shown to have significant heritabilities for larval growth. Secondly, we estimated the lipid content of occytes from female parents, since this is hypothesized to affect larval growth and survival. Finally, we estimated the among-family variation in mortality from 4 to 21 d as a test of possible genetic variation for viability among larval scallops. Significant genetic variation (h²=1.10 to 1.24) was estimated for larval shell length at 4, 14, and 21 d. There was a significant correlation (r=0.66) between larval size at 4 d and lipid content of oocytes, but only when two females with high levels of lipid oocyte^–1 were excluded as outliers. There was no correlation between larval size at 14 and 21 d and lipid content of oocytes. Mortality among families from 4 to 21 d was high (69 to 97%), and was significantly different among families. These results indicate that there is significant heritability for larval growth which is largely independent of the lipid content of the oocytes. A high heritability for larval growth may indicate that this trait is only weakly correlated with fitness.     

Threat-sensitive foraging by larval threespine sticklebacks (Gasterosteus aculeatus)

Summary The threat-sensitive predator avoidance hypothesis predicts that prey can assess the relative threat posed by a predator and adjust their behaviour to reflect the magnitude of the threat. We tested the ability of larval threespine sticklebacks to adjust their foraging in the presence of predators by exposing them to conspecific predators of various sizes and recording their foraging and predator avoidance behaviours. Larvae (<30 days post-hatch) displayed predator escape behaviours only towards attacking predators. At 3 weeks post-hatch larvae approached the predator after fleeing, a behaviour which may be the precursor to predator inspection. Larvae reduced foraging and spent less time in the proximity of large and medium-sized predators compared to small predators. The reduction in foraging was negatively correlated to the predator/larva size ratio, indicating that larvae increased their foraging as they increased in size relative to the predator. We conclude that larval sticklebacks can assess the threat of predation early in their ontogeny and adjust their behaviour accordingly.Correspondence to: J.A. Brown     

Calcification of the first larval shell of Tridacna squamosa (Tridacnidae: Bivalvia)

The transistional stage between the trochophore and the veliger larva of Tridacna squamosa Lamarck, including shell formation and initial organogenesis, is described. Calcification of the Prodissoconch I begins before formation of the organic component of the shell is completed, and proceeds in a pattern which reinforces the maximum stress areas of the shell, allowing the early onset of the larval escape behavior. Evidence is presented which suggests that the orientation of the mineral crystals in the larval shells is determined by an organic component of the shell and not by an active process of the clam during calcification.     

Functional morphology of the larval tentacles of Phragmatopoma californica (Polychaeta: Sabellariidae): composite larval and adult organs of multifunctional significance

Larvae of the sabellariid polychaete Phragmatopoma californica (Fewkes), which were collected in San Diego, California and were competent to metamorphose after 18 to 30 d of development were observed in vivo by videoequipped light microscopy, and the fine structure of the larval tentacles was examined by transmission and scanning electron microscopy. Each tentacle has tufts of at least two types of immotile cilia arranged in dorsolateral and ventrolateral rows, and a ventral groove covered by two types of motile cilia that beat independently of each other. The epidermis is regionalized into glandular, sensory, locomotory, and support cell types and contains four longitudinal bundles of basiepithelial nerve fibers. A layer of connective tissue separates the epidermis and the nerve tracts from obliquely striated muscles that occur within the peritoneum that lines the central coelomic cavities. The peritoneum forms an intact coelomic epithelium that overlies and interdigitates with the muscle cells, with no intervening basal lamina. The muscle cells are considered to be intraperitoneal because they are located above the basal lamina and they lack intercellular junctions with the peritoneal cells. Specialized peritoneal cells form a striated myoepithelial blood vessel that partitions the coelom into medial and lateral cavities. No neuromuscular junctions were found, but both muscular and ciliary movement seem to be under neuronal control. The basiepithelial nerve terminals appear to synapse into the connective tissue layer toward the intraperitoneal muscle. Several similarities in tissue organization are noted between the larval tentacles of P. californica and the tube feet of echinoderms. Observations on the ontogeny, morphology, and behavior of the tentacles suggest that they are multifunctional organs involved in feeding, construction of the juvenile sand tube, locomotion, attachment, and sensory perception during larval and adult lives.     

Evolution of reproductive features and larval development in the genus Amphiglena (Polychaeta: Sabellidae)

Larval development in Amphiglena mediterranea is direct in the parental tube, and larvae crawl away as young juveniles with a radiolar crown capable of feeding. The non-feeding larvae of A. nathae are initially brooded in the parental tube, and then have a swimming phase before settling and developing a radiolar crown. Developmental data and reproductive features such as sperm structure are combined with other morphological data for an assessment of the cladistic relationships of the species in the genus Amphiglena. Two species of Laonome are included as part of the ingroup. Six genera, representing a sister group to the Laonome/Amphiglena clade, are used as an outgroup. Two most-parsimonious cladograms were found, and the evolution of reproductive features are discussed. The monophyly of Amphiglena is indicated by features previously identified, as well as two features based on sperm morphology. At present a plesiomorphic larval form cannot be identified for the genus Amphiglena. Received: 16 June 1997 / Accepted: 29 January 1998