Effects of a transitory,low food supply in the early life of larval herring (Clupea harengus) on mortality,growth and digestive capacity

In order to assess possible effects of a transitory, low food supply on later development, three groups of Clyde herring larvae (Clupea harengus L.) were exposed in 1989 to different feeding regimes immediately after yolk resorption. Group 1 received a high daily ration of 80 copepods larvae^–1 for 31 d, Group 2 a low daily ration of 15 copepods larva^–1 for 10 d followed by a high ration (80 copepods larva^–1) for 21 d and Group 3 a low ration of 15 to 20 copepods larva^–1 for 31 d. After 31 d of feeding, digestive capacity, expressed as the sum of trypsin and trypsinogen, was markedly reduced in Group 2 compared to Group 1, while Group 3 had an even lower digestive capacity. After the switch from low to high ration Group 2 exhibited compensatory growth and caught up with Group 1 both in standard length and content of soluble protein. Group 3 had the lowest growth rates. Mortality was equal in Groups 1 and 2, while Group 3 showed an excess mortality of 40% of the start population. Although Group 2 larvae had caught up with Group 1 in growth at the end of the study, content of trypsin and trypsinogen in Group 2 was only half of that found in Group 1. Thus, comparing effects of a short period of food limitation on future growth, mortality and content of digestive enzymes, the study indicates content of trypsin and trypsinogen to be the most sensitive variable for detection of food limitation in the early stages of exogenous feeding.     

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.     

Osmoacclimation in Enteromorpha intestinalis: long-term effects of osmotic stress on organic solute accumulation

The fate of the protease trypsin in intestines of individual herring larvae Clupea harengus L. was studied following digestion of the copepod Acartia tonsa. Trypsin was retained in the intestine during two consecutive pulses of feeding and defaecation of copepods. Quantification of herring trypsin in digested, defaecated copepods showed that ca. 1% of larval intestinal enzyme was defaecated along with 1 to 3 copepods. Following ingestion of a single meal, the level of intestinal trypsin post-ingestion declined to pre-ingestion levels within 1 to 2 d of starvation. All enzyme data thus indicated that trypsin, released in response to ingestion of a meal, was retained. In addition, analysis of fed subgroups of starved larvae clearly indicated that release of trypsin from the pancreas stopped after 6 to 8 d of starvation. As the fish still contained substantial amounts of trypsinogen, the underlying cause might be defective release mechanisms. Daily secretion of trypsin and processes responsible for enzyme retention in the gut are discussed. Assimilation efficiency in herring larvae was estimated for copepodite prey. Average carbon assimilation was 90%.     

Induction of trypsinogen secretion in herring larvae (Clupea harengus)

Mechanisms initiating trypsinogen secretion were studied in laboratory reared herring larvae (Clupea harengus L.) exposed to physical and chemical stimuli. Pancreatic secretion of trypsinogen was quantified for each stimulus type as the increase above pre-stimulus level of intestinal trypsin content. Larval prey types were: nauplii, copepodites or adult Acartia tonsa, small polystyrene spheres (diameter 94 m), small (diameter 79 m) or large (diameter 170 m) polystyrene-latex spheres. Intestinal trypsin content can be expressed as a function of two variables: meal size and content of pancreatic trypsinogen. Trypsinogen secretion increases with different prey items in the order: small spheres, nauplii and copepodites. Larvae which eat large spheres secrete more enzyme than if fed small spheres but trypsinogen secretion is similar in fish larvae fed copepodites and large spheres. The fact that the size of non-biodegradable particles exerts a major control over trypsinogen secretion suggests neural — as opposed to chemically mediated — initiation of secretion. A cephalic phase of secretory stimulation could not be demonstrated during swallowing of copepods or exposure for 2 to 3 h to compounds which leak from live copepodites. As cephalic and gastric phases of secretory stimulation are absent, initiation of trypsinogen secretion must take place in the intestine. Larval herring retain trypsin in the intestine. Ca. 4.5 h after a meal, 3/4 of the enzyme is located in the intestinal fluid, presumably available for hydrolysis of subsequent meals, and the high proportion (ca. 25%) of the pancreatic trypsinogen content which is secreted for copepodite prey may thus not be energetically wasteful for the larvae.     

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.     

Food limitation stimulates metamorphosis of competent larvae and alters postmetamorphic growth rate in the marine prosobranch gastropodCrepidula fornicata

The effects of food limitation on growth rates and survival of marine invertebrate larvae have been studied for many years. Far less is known about how food limitation during the larval stage influences length of larval life or postmetamorphic performance. This paper documents the effects of food limitation during larval development (1) on how long the larvae ofCrepidula fornicata (L.) can delay metamorphosis in the laboratory after they have become competent to metamorphose and (2) on postmetamorphic growth rate. To assess the magnitude of nutritional stress imposed by different food concentrations, we measured growth rates (as changes in shell length and ash-free dry weight) for larvae reared in either 0.45-m filtered seawater or at phytoplankton concentrations (Isoehrysis galbana, clone T-ISO) of 1 × l0³, 1 × 10⁴, or 1.8 × 10⁵ cells ml^–1. Larvae increased both shell length and biomass at 1 × 10⁴ cells ml^–1, although significantly more slowly than at the highest food concentration. Larvae did not significantly increase (p > 0.10) mean shell length in filtered seawater or at a phytoplankton concentration of only 1 × 10³ cells ml^–1, and in fact lost weight under these conditions. To assess the influence of food limitation on the ability of competent individuals to postpone metamorphosis, larvae were first reared to metamorphic competence on a high food concentration ofI. galbana (1.8 × 10⁵ cells ml^–1). When at least 80% of subsampled larvae were competent to metamorphose, as assessed by the numbers of indlviduals metamorphosing in response to elevated K⁺ concentration in seawater, remaining larvae were transferred either to 0.45-m filtered seawater or to suspensions of reduced phytoplankton concentration (1 × 10³, 1 × 10⁴, or 5 × 10⁴ cells ml^–1), or were maintained at 1.8 × 10⁵ cells ml^–1. All larvae were monitored daily for metamorphosis. Individuals that metamorphosed in each food treatment were transferred to high ration conditions (1.8 × 10⁵ tells ml^–1) for four additional days to monitor postmetamorphic growth. Competent larvae responded to all food-limiting conditions by metamorphosing precociously, typically 1 wk or more before larvae metamorphosed when maintained at the highest food ration. Surprisingly, juveniles reared at full ration grew more slowly if they had spent 2 or 3 d under food-limiting conditions as competent larvae. The data show that a rapid decline in phytoplankton concentration during the larval development ofC. fornicata stimulates metamorphosis, foreshortening the larval dispersal period, and may also reduce the ability of postmetamorphic individuals to grow rapidly even when food concentrations increase.     

The effect of food availability,age or size on the RNA/DNA ratio of individually measured herring larvae: laboratory calibration

RNA/DNA ratios in individual herring (Clupea harengus) larvae (collected from Kiel Bay, Baltic Sea, in 1989) were measured and proved suitable for determining nutritional status. Significant differences between fed and starving larvae appeared after 3 to 4 d of food deprivation in larvae older than 10 d after hatching. The RNA/DNA ratio showed an increase with age or length of the larvae and was less pronounced in starving larvae compared to fed larvae. The individual variability of RNA/DNA ratios in relation to larval length of fed larvae and of larvae deprived of food for intervals of 6 to 9 d is presented. Based on the length dependency and the individual variability found within the RNA/DNA ratios, a laboratory calibration is given to determine whether a larva caught in the field has been starving or not. An example for a field application is shown.     

Differential timing of larval starvation effects on filtration rate and growth in juvenile Crepidula onyx

This study demonstrates that the timing of larval starvation did not only determine the larval quality (shell length, lipid content, and RNA:DNA ratio) and the juvenile performance (growth and filtration rates), but also determine how the latent effects of larval starvation were mediated in Crepidula onyx. The juveniles developed from larvae that had experienced starvation in the first two days of larval life had reduced growth and lower filtration rates than those developed from larvae that had not been starved. Lower filtration rates explained the observed latent effects of early larval starvation on reduced juvenile growth. Starvation late in larval life caused a reduction in shell length, lipid content, and RNA:DNA ratio of larvae at metamorphosis; juveniles developed from these larvae performed poorly in terms of growth in shell length and total organic carbon content because of “depletion of energy reserves” at metamorphosis. Results of this study indicate that even exposure to the same kind of larval stress (starvation) for the same period of time (2 days) can cause different juvenile responses through different mechanisms if larvae are exposed to the stress at different stages of the larval life.     

Temporal and spatial variation in the growth rates of Baltic herring (<Emphasis Type="Italic">Clupea harengus membras</Emphasis> L.) larvae during summer

It has been suggested that larval survival determines the year-class strength in most marine fish species. During their growth and development, the ability of the larvae to catch prey and avoid predation will increase. However, the factors affecting short-term changes in the growth of Baltic Sea herring have been little studied in the field. We collected Baltic herring (Clupea harengus membras L.) larvae from five different towing areas in the Archipelago Sea (SW Finland) during May and June 1989, right after the main spawning season. Twenty thousand two hundred and ten larvae were analysed and the area-specific growth rate (i.e. increase in standard length) was estimated by tracing the larval cohorts from the length-frequency data. This represents the first Baltic herring study with daily sampling during a long study period. The growth rate was related to environmental factors, such as temperature, number of zooplankters, and wind speed and direction. Large variation in larval growth rate occurred between areas: lowest and highest growth rates were 0.18 and 0.52 mm·day^-1. Temperature was an important variable controlling larval-fish growth rate. An increase of one 1°C in average water temperature corresponded to an increase in growth rate of 0.043 mm·day^-1. This may have been caused either by a direct temperature effect (changes in metabolic rate) or by the indirect effect of changes in food availability. We also found the densest herring populations in the areas with highest average water temperature. However, temperature and larval growth rate both increased towards the inner archipelago.     

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.     

Effects of algal and larval densities on development and survival of asteroid larvae

Effects of larval and algal culture density and diet composition on development and survival of temperate asteroid larvae were studied in the laboratory at Santa Cruz, California, USA, during summer and fall of 1990. Larvae of Asterina miniata were reared at two densities, 0.5 or 1.0 ml^-1, and fed one or two species of cultured phytoflagellates — Dunaliella tertiolecta alone or mixed with Rhodomonas sp. — at three concentrations of 5x10², 5x10³, and 5x10⁴ total cells ml^-1. Algal concentration strongly influenced larval development; however, larval density also had a marked effect. Development progressed further with increasing algal concentration. Larval growth and differentiation were sometimes uncoupled; i.e., growth measures were directly related to food level, while differentiation indicators were less so. At the lowest food level, growth was negative and differentiation was arrested at early precompetent stages; these larvae never formed juvenile rudiments or brachiolar attachment structures. Development times of larvae given more food ranged from 26 to 50 d and depended directly on food availability. Development time to metamorphosis at the highest food concentration was similar for siblings fed D. tertiolecta alone or mixed with Rhodomonas sp. In contrast, when food level was an order of magnitude lower, larvae fed the algal mixture metamorphosed significantly earlier than larvae fed the unialgal diet. This suggests interactive effects of food quantity and food quality. Survival was little affected by larval or food density, except at the lowest ration. Feeding experiments in well-controlled laboratory conditions are useful to predict and compare the physiological or developmental scope of response of larvae to defined environmental factors; however, results from such studies should not be extrapolated to predict rates and processes of larval development in nature.     

Trypsin content in intestines of herring larvae,Clupea harengus,ingesting inert polystyrene spheres or live crustacea prey

In 1986, at the Danish Institute of Fisheries and Marine Research, Denmark, Clupea harengus L. larvae from three different herring stocks were offered either non-biodegradable polystyrene spheres, nauplii and copepodites of Acartia tonsa or Artemia ssp. nauplii. Ingestion of polystyrene spheres induced trypsin secretion to a higher level than in non-feeding fish. Larvae ingesting live food of the same width as the polystyrene spheres exhibited the highest trypsin content in the intestines. Mechanisms responsible for the regulation of pancreatic enzyme secretion are discussed.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

Factors influencing predation of Hyperoche medusarum (Hyperiida: Amphipoda) on larvae of the Pacific herring Clupea harengus pallasi

Predation of different-sized Hyperoche medusarum (Hyperiida: Amphipoda) on larvae of the Pacific herring Clupea harengus pallasi was studied in the laboratory. The attacking rate of H. medusarum was a function of herring larvae size as well as size of the predator, and varied from 0.15 to 0.95 larvae attacked h^-1 per hyperiid. In the range of 7.55 to 16.05 mm total larval length, vulnerability to predation was highest for 13.3 and 13.7 mm larvae. Large hyperiids swam faster and covered a wider area during searching and were more effective predators than small ones. Predation seemed to be influenced by light, and its intensity was dependent on the duration of previous food deprivation of the hyperiid.This study was sponsored by the International Bureau of the Gesellschaft für Kernenergiever-wertung in Schiffbau und Schiffahrt in connection with the German Canadian agreement on scientific and technical cooperation.     

Effects of delayed feeding and temperature on the age of irreversible starvation and on the rates of growth and mortality of Pacific herring larvae

The time periods from exhausion of the yolk to the age of irreversible starvation for Pacific herring Clupea harengus pallasi larvae were 8.5, 7.0 and 6.0 d at 6°, 8° and 10°C, respectively. These periods are within the range perviously measured for Atlantic herring larvae and other temperature zone fish species; they are long compared to the periods for tropical species. The variation in the length of this period is due almost entirely to temperature; the natural logarithm of the time period from fertilization to irreversible starvation is highly correlated (r=0.91) with the mean rearing temperature for 25 species of pelagic marine fish larvae. The rates of growth and mortality, measured for 26 experimental populations of Pacific herring larvae reared at 6°, 8° and 10°C and ten ages of delayed first feeding, decreased and increased, respectively with increasing age of first feeding and increasing temperature. These rates, adjusted for the effects of rearing conditions, were compared with the rates for natural populations of herring larvae. Growth is generally faster in the sea than in experimental enclosures. Two of the eleven estimates of natural mortality rate were high enough to indicate possible catastrophic mass starvation. This is consistent with Hjort's critical period concept of year class formation and it suggests that mass starvation occurs in 18 to 36% of the natural populations of first feeding herring larvae.     

Ontogenetic changes in the vertical distribution of giant scallop larvae,Placopecten magellanicus,in 9-m deep mesocosms as a function of light,food, and temperature stratification

To understand how thermal stratification and food abundance affects the vertical distribution of giant scallop larvae Placopecten magellanicus (Gmelin), a mesocosm study was conducted in January and February 1992. The position of larvae was followed over 55 d in replicated 9-m deep tanks in relation to a sharp thermocline and the presence or absence of phytoplankton. Growth and vertical position of larvae were monitored in separate treatments which included phytoplankton added above the thermocline, below the thermocline, throughout the mesocosm, or absent from the mesocosm. Changes in the vertical position of larvae over time were quantified with a new, profiling, video-optical instrument capable of semi-automatically identifying, counting and sizing larvae. The strong diurnal migration of scallop larvae resulted in aggregations at two interfaces: the air/water interface during the night, and at the thermocline during the day. At times, the concentration of larvae within cm of the surface was > 100 times that in the remaining water column. The formation of bioconvective cells of swimming larvae at the air/water interface allowed larval aggregations to persist throughout the period of darkness. Regardless of the distribution of food, larvae remained above the thermocline during most of the experiment. Therefore, only in those treatments where food was also present above the thermocline was larval growth relatively high. Larger larvae penetrated the thermocline only after reaching a shell length of about 200 m; thus larval size, rather than chronological age, was more important in describing their vertical distribution. The rapid increase in kinematic viscosity with decreasing water temperature at the thermocline may retard the movement of larvae and contribute to aggregation at this interface. The influence of larval size on their vertical distribution, and the resulting potential for horizontal transport to settlement sites, points to the importance of persistent hydrographic features as critical factors contributing to settlement variance in scallops.     

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.     

Morphological and histological changes during the growth and starvation of herring and plaice larvae

Reared herring (Clupea harengus L.) and plaice (Pleuronectes platessa L.) were examined for morphological and histological changes during growth and starvation. The growth rate of herring larvae of 0.22 mm/day was less than that reported for wild stock, but this difference was attributed to survival of runts in laboratory. Larval plaice had a growth rate of 0.16 mm/day. The relative condition factor (antilogarithm of intercept of length-weight line) was used to assess condition throughout the larval stages. Starvation resulted in a progressive collapse of the larval body, especially of the ventral body surface around the pectoral girdle of both species (assessed by the pectoral angle) and of the spacing between the organs of the head in herring. There was a breakdown of the herring gut with decreases in epithelial cell height and catabolism of the connective tissue coat and a marked reduction in the transverse sectional area of the plaice liver. The changes in the pectoral angle in both herring and plaice and the eye height to head height ratio in herring should be useful to fishery biologists for assessing nutritional condition, even on board ship.     

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.     

Effects of temporary starvation on larvae of the sea star Asterina miniata

Patchy food distribution may force temporary starvation conditions on planktonic larvae. This potential food limitation may affect survivorship, duration of larval period, and post-metamorphic succes. In this study, larvae of the asteroid Asterina miniata were subjected to temporary food deprivation of several durations and at different stages. Developmental effects were documented by quantification of larval stage, total length, time to metamorphosis, initial juvenile radius, range of settling times, and percent survival to metamorphosis. All starved treatments were significantly affected in settling time and most in percent survival. However, larvae starved later in development demonstrated tremendous tolerance of food deprivation (e.g. the total number of settlers in the treatment starved for 28 d was not significantly different from the fed control). Survival was lower in treatments starved earlier in development than those starved later. Food is apparently required until late in larval development to facilitate metamorphosis. The range of settling times was large; for example, the continuously-fed control treatment produced juveniles from Days 58 through 136. Temporary starvation had no effect on initial juvenile radius.