Energy content at metamorphosis and growth rate of the early juvenile barnacle<Emphasis Type="Italic"> Balanus amphitrite</Emphasis>

The energetic cost of metamorphosis in cyprids of the barnacle Balanus amphitrite Darwin was estimated by quantification of lipid, carbohydrate and protein contents. About 38–58% (4–5 mJ individual^–1) of cypris energy reserves were used during metamorphosis. Lipids accounted for 55–65%, proteins for 34–44% and carbohydrates for <2% of the energy used. Juveniles obtained from larvae fed 10⁶ cells ml^–1 of Chaetoceros gracilis were bigger (carapace length: 560–616 µm) and contained more energy (5.56±0.10 mJ juvenile^–1) than their counterparts (carapace length: 420–462 µm; energy content: 2.49±0.20 mJ juvenile^–1) obtained from larvae fed 10⁴ cells ml^–1. At water temperatures of 30°C and 24°C and food concentrations of 10⁴ and 10² cells ml^–1 (3:1 mixture of C. gracilis and Isochrysis galbana) as well as under field conditions (26.9±3.1°C and 2.2±0.8 µg chlorophyll a l^–1), juveniles obtained from larvae fed the high food concentration grew faster than juveniles obtained from larvae fed low food concentration until 5 days post-metamorphosis. Laboratory experiments revealed a combined effect of early juvenile energy content, temperature and food concentration on growth until 5 days post-metamorphosis. After 10 days post-metamorphosis, the influence of the early juvenile energy content on growth became negligible. Overall, our results indicate that the energy content at metamorphosis is of critical importance for initial growth of juvenile barnacles and emphasize the dependency of the physiological performance of early juvenile barnacles on the larval exposure to food.Communicated by O. Kinne, Oldendorf/LuheAn erratum to this article can be found at     

Growth of sea bass,Dicentrarchus labrax,larval and juvenile stages and their otoliths under quasi-steady temperature conditions

The relationship between somatic growth and growth of otoliths of sea bass larvae, postlarvae and juveniles under relatively steady temperature conditions was studied. Larvae were incubated at the constant ambient temperature of 13.5°C, whereas postlarvae and juveniles were reared at a comparatively steady temperature ranging from 18.6 to 20.4°C, with a mean of 19.67°C. The patterns of both somatic and otolith growth were found to be similar. Differentiated data on larvae length and otolith diameters indicated three periods of change in their growth rates. Since temperature was kept relatively steady during the experiment, and larvae fed ad libitum, these periods could be attributed with relative certainty to intrinsic changes which occur during stage-specific periods of growth. The third period of change in both growth rates indicates a specific phase of growth during metamorphosis. The changes in growth rates, as well as the raw time series of the growth of both larval lengths and otolith diameters, may be described by higher order polynomials with a high degree of probability levels. A non-linear relationship between body length and otolith diameters was established, indicating positive allometric growth of otoliths. It was also observed that the coefficient of allometric growth changed at the time estimated for the end of metamorphosis. Thus, a non-linear relationship and changes in the coefficients of allometry should be borne in mind when back-calculating somatic growth from the growth of otoliths.     

Physiological and biochemical changes during lecithotrophic larval development and early juvenile growth in the northern stone crab,Lithodes maja (Decapoda: Anomura)

Larvae of the northern stone crab, Lithodes maja L., were reared in the laboratory from hatching to the second crab stage. complete larval development (at constant 9°C) lasted about 7 wk, invariably consisting of three pelagic zoeal stages and a semibenthic Megalopa; only two zoeal stages have been described in the literature. All larval stages are lecithotrophic. First feeding was consistently observed only after metamorphosis, in the first juvenile crab stage. In short intervals (every 1 to 5 d), developmental changes in biomass, B (expressed as: dry weight, W; carbon, C; nitrogen, N; hydrogen, H) and oxygen consumption (respiration, R) were measured in larvae and early juveniles; additionally, protein and carbohydrates were measured, but only in the zoeal stages and early Megalopa. Unusually high C contents (varying between 56 and 61% of W in eggs and freshly hatched Zoea I larvae from 12 different females) and high C:N weight ratios (8 to 11) indicate enhanced initial lipid stores, which are utilized as the major metabolic substrate during both embryonic and lecithotrophic larval development. Predominant degradation of lipids is shown indirectly; the C:N ratio decreased significantly, from 10 (at hatching) to 6 (at metamorphosis), while larval protein decreased only little, from ca. 55% of W (at hatching) to 48% (in the Megalopa). From hatching to metamorphosis, about 27% of the initially present W, 48% of C, 18% of N, and 52% of H were lost. This decrease in larval biomass can be described as an exponential function of development time. The major part of these losses were associated with metabolic energy requirements, while exuvial losses were comparably small. In each of the zoeal stages, only about 1 to 2% of late premoult (LPM) B was shed with the exuvia. The Megalopa, which produces a much thicker, calcified exoskeleton, lost 20% of LPM W, but only 5 to 8% of organic constituents (C, N, H). Much higher exuvial losses were measured in the Crab I stage (51% in W, 21% in C, 5% in N, and 7% in H). Maximum respiration was found in the actively swimming zoeal stages, a minimum in the predominantly benthic, mostly inactive Megalopa. The Crab I stage exhibits also a sluggish behaviour and low R, in spite of beginning food uptake and growth. Immediately after metamorphosis, the juvenile crab gained rapidly in W, in particular in its C fraction. A transitorily steep increase in the C:N ratio indicates a replenishment of partially depleted lipid stores, but also a rapid initial increase of inorganic C in the heavily calcified exoskeleton. Instantaneous rates of growth, assimilation, and net growth efficiency (K ₂) were high during the initial (postmoult) phase in the first juvenile crab stage (C-specific growth rate: 6% d^-1; K ₂:70%), but decreased towards zero values during laterstages of the moulting cycle; metabolism remained practically constant during the Crab I stage. Entirely lecithotrophic larval development from hatching to metamorphosis in L. maja is considered an adaptation to seasonally short and limited planktonic food production in subarctic regions of the northern Atlantic.     

Growth and metamorphosis of Aplysia oculifera larvae in laboratory culture

Eggs of Aplysia oculifera (Adams and Reeve, 1850) were incubated in the laboratory. They hatched 8 to 9 d after spawning. Shell length (SL) of the hatched larvae was 102±2 m. Larvae were fed on the unicellular algae Isochrysis galbana in a concentration of 10⁴ cell ml^-1, and after 45 to 60 d grew to a maximum SL of 385±11 m. Larvae survived up to 330 d. A total of 12 species of algae from the natural habitat of A. oculifera were examined as metamorphosis inducers. Red algae Dasia sp., Jania sp., Hypnea sp. and Liagora sp. induced metamorphosis in 66.7±21.2, 28.3±17.7, 26.0±18.5 and 4.0±8.0% of the larvae, respectively. Green algae Enteromorpha intestinalis and Ulva sp. induced metamorphosis in 37.0±11.0 and 9.0±10.4% of the larvae, respectively. Cladophora sp. and Codium dichotomum, and the brown algae Padina pavonia, Colpomenia sinuosa, Hydroclathrus clathratus and Cystoseira sp. did not induce metamorphosis. There was no significant difference in the rate of metamorphosis between young (2 to 4 mo) and old (6 to 8 mo) larvae. Postmetamorphic juveniles grew and developed only when fed with E. intestinalis. They grew to a body length of>8 mm in 50 d. Postmetamorphic juveniles did not survive on other algae. The longevity of the planktonic A. oculifera larvae supports the hypothesis that the larvae can exist in the plankton and survive for several months until the next recruitment. The advantage of non-specificity in metamorphosis induction is discussed.     

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.     

Sub-lethal impact of carbaryl on food utilization in the freshwater prawn Macrobrachium malcolmsonii

This study determines the toxic effect of carbaryl (Sevin50% W.P) on the food utilization parameters in intermoult juveniles of the prawn, Macrobrachium malcolmsonii. The prawns (4.5-5.0 cm in length and 1.0-1.25 g wet wt.) were exposed to three sub-lethal concentrations of carbaryl (5.15, 7.73 and 15.47 microgl-1) for duration of 40 days. The toxic medium was renewed daily. The prawns were fed ad libitum with known energy quantity of boiled goat liver on daily basis. The overall wet weight gain was calculated. The energy lost through unconsumed food (15-60%), faeces (15-109%), ammonia excretion (9-27%) and moults (13-26%) of the prawns were calculated. The feeding rate, the rate and efficiency of absorption, the metabolic and food conversion rates and the gross and net food conversions efficiencies were found to be significantly declined (p<0.05) in test prawns when compared to that of the control. The energy lost through faeces, ammonia excretion and exuvia was found to be significantly elevated (p<0.05) in test prawns than that of the control. The effectof carbaryl on the bioenergetics parameters was severe in the highest sub-lethal concentration, less in the intermediate concentration and least in the lowest sub-lethal concentration. The results indicated that decrease in feeding, absorption, metabolism and food conversion are interdependent and toxicity of carbaryl diverting energy from production to maintenance pathways, which ultimately resulting in declined growth of M. malcolmsonii.     

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.     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.     

The development of functional digestive and metabolic organs in turbot,Scophthalmus maximus

The functional status of organ systems involved into the processing of exogenous food is critical for the survival and growth of fish early life stages. The present study on laboratory-reared larval turbot, Scophthalmus maximus, provides an overview on the ontogeny of structure and functions involved in digestion, absorption and metabolism of nutrients. At start of exogenous feeding the intestine of larval turbot is anatomically differentiated, with enterocytes displaying an adult-type ultrastructure and being able to process lipids. At the microvillous border of the enterocytes, enzymes of contact digestion such as aminopeptidase are found. The ultrastructure of the exocrine pancreatic cells is fully differentiated from hatching onwards. Likewise, substantial activities of trypsin-type proteases are present. A stomach anlage exists in first-feeding larvae; however, the stomach becomes functional (appearance of gastric glands and pepsin secretion) only during metamorphosis. Liver parenchymal cells already display a functional ultrastructure during the endotrophic phase; with onset of exogenous feeding they develop pronounced diet-related changes of their energy stores. Larval respiration is not executed by the gills since respiratory surface of these structures develops only towards metamorphosis. The energy generation of larval muscle tissue depends on aerobic metabolism, whereas glycolytic activities start to increase at metamorphosis. In conclusion, two important patterns can be recognized in the development of turbot larvae: (1) The structure/function is differentiated at hatching or at the onset of exogenous feeding (afterwards it experiences mainly quantitative but not qualitative growth, i.e., intestine, exocrine pancreas, liver); or (2) the structure/function is absent in larvae and develops only during metamorphosis (i.e., gills, glycolytic muscle metabolism, stomach).     

Role of biochemical energy reserves in the metamorphosis and early juvenile development of the oyster Ostrea chilensis

Metamorphosis in the Chilean oyster Ostrea chilensis was complete 36 h after release of the larvae, when 100% of the individuals showed edge growth of the dissoconch. The size of the larval shell did not change during metamorphosis, although the total dry weight of the larva decreased considerably. During this period, when the gill ciliature was undeveloped and the oyster therefore unable to feed, energy demands were met by biochemical reserves retained from the larval phase. Proteins contributed the largest quantity of energy to the metamorphosing oyster, 69.3% of the total expended, whereas lipids supplied 24.3% and carbohydrates only 6.4%. The process of metamorphosis consumed 64.5% of the energy reserves held by the pediveliger at the time of release. When metamorphosis was complete, growth began and tissue reserves were replenished, protein and carbohydrate accumulating rapidly early in the juvenile stage. Received: 26 December 1997 / Accepted: 8 July 1998     

Energetic cost of development through metamorphosis for the seastar<Emphasis Type="Italic"> Mediaster aequalis</Emphasis> (Stimpson)

Rates of oxygen consumption were measured for embryos, larvae and juveniles of the seastar Mediaster aequalis for 76 days post-fertilization. The rate increased from 0.65 nmol O₂ ind^–1 h^–1 at 6 h after fertilization to 2.8 nmol O₂ ind^–1 h^–1 at day 35. Larvae became competent to metamorphose around day 35 post-fertilization and began to decrease their metabolic rate after this time. Metamorphosed juveniles consumed 0.74 nmol O₂ ind^–1 h^–1. Eggs contained 138.6 µg lipid ind^–1 and 12.1 µg protein ind^–1. Lipid levels decreased in concentration throughout development while protein levels increased slightly. The lipid levels decreased by 88.5 µg from eggs to day 76 larvae, accounting for 3.5 J of energy. Total oxygen consumption to this point was 3.74 µmol O₂ ind^–1, accounting for 1.84 J. The energetic demand up to day 76 was met completely through the use of lipid reserves. Metamorphosed juveniles expended 0.5 J more than larvae at the same age. Tubes of the polychaete Phyllochaetopterus prolifica were able to induce metamorphosis in M. aequalis larvae and a non-polar extract of these tubes also triggered metamorphosis. Larvae that are delayed to metamorphose can sustain their metabolic rate with lipid reserves for a limited, yet undetermined, period.Communicated by P.W. Sammarco, Chauvin     

Leptocephalus energetics: assembly of the energetics equation

The principles of energetics were used to examine the energetic requirements of leptocephali. Respiration and excretion rates and daily growth rates combined with proximate composition were used to examine the allocation of energy into each of the three main components of energetics: metabolism, excretion and growth. The daily energetic requirements for leptocephali, referred to as type 2 larvae based upon their unique developmental strategy, were compared to the requirements of non-leptocephalus larvae, known as type 1. Leptocephalus daily energetic requirements were also compared to the energy available from the leptocephalus' proposed food sources. The four species of eel larvae selected were all from the order Anguilliformes: Paraconger caudilimbatus (Poey), Ariosoma balearicum (Delaroche), Gymnothorax saxicola Jordan and Davis, and Ophichthus gomesii (Castelnau). The allocation of energy to each of the components of energetics as well as the total energetic requirements for the leptocephali proved to be very different from those of type 1 larvae. Metabolism received the majority, 60-92%, of the energy required per day. Growth and excretion were allocated 4-39% and <1-21%, respectively, of the total energy needed per day. Leptocephali required <50% of the energy needed by type 1 larvae of equal dry mass. The unique growth strategy used by leptocephali allows them to increase rapidly in size while allocating the majority of their energy, not to growth as in most larval fish, but to metabolism.     

Histological study of the effects of starvation on reared and wild-caught larval stone flounder,Kareius bicoloratus

Starvation tolerance of laboratory-reared larval stone flounder, Kareius bicoloratus, was examined at different temperatures and salinities during the winters of 1984, 1985 and 1986. Starvation tolerance decreased with increased temperature and exhibited low values with high salinities. The highest starvation tolerance observed at low salinity was just before the metamorphosis stage. Starvation tolerance showed little change until larvae were 11 d. It increased with age thereafter. Epithelial cell heights of the digestive tract and cell diameters of pancreas and liver were measured histologically in reared stone flounder during growth and starvation. These values decreased markedly in the starved condition. Aldehyde fuchsin positive granules in the rectal epithelium also disappeared during the short starved period. The nutitrional condition of wild-caught stone flounder larvae, collected in January and February 1986 from the Matsukawa-ura inlet, Fukushima, Japan, was also examined. Eighty percent of larvae were estimated to be in fed condition just before sampling. The changes in cell heights of digestive organs agreed with this estimate. These histological methods seem to be useful in assessing the nutritional condition of marine fish larvae.     

Use of kappa-carrageenan microbound diet (C-MBD) as feed for Penaeus monodon larvae

The performance of an artificial practical diet, kappacarrageenan microbound diet (C-MBD) was assessed on Penaeus monodon larvae at the SEAFDEC Broodstock and Maturation Experimental Laboratory in March 1986. Shrimps were reared from zoea₁ to post-larvae₁ using five dietary treatments: (a) natural food — Chaetoceros calicitrans and Artemia salina (b) C-MBD; (c) combination of natural food and C-MBD; (d) commercial diet (microencapsulated, MED); (e) combination of natural food and commercial diet. Results showed slow development with larvae fed the commercial diet. Feeding with C-MBD in combination with natural food resulted in the highest % survival among treatments (69.6), but this was not significantly different (P>0.05) from those obtained with larvae fed natural food alone, C-MBD alone or their combination. While mean values for survival of larvae fed the commercial diet, either alone or in combination, was significantly lower (p<0.05) than all other treatments, their mean growth indices were comparable with larvae fed C-MBD alone or in combination. The low levels of protein, lipid and essentially fatty acids (which are considered important nutrients during larval development) contained in the commercial diet may well justify the results on metamorphosis, survival and growth of the larvae fed this diet. The good performance of C-MBD in this experiment suggests that this kind of diet can be used as partial or total replacement to the traditional algal food.     

Growth and differentiation during delayed metamorphosis of feeding gastropod larvae: signatures of ancestry and innovation

Extent of larval growth among marine invertebrates has potentially profound implications for performance by benthic recruits because body size influences many biological processes. Among gastropods, feeding larvae often attain larger size at metamorphic competence than non-feeding larvae of basal gastropod clades. Delay of metamorphosis can further influence size at recruitment if larvae continue to grow during the delay. Some caenogastopod larvae grow during delayed metamorphosis, but opisthobranch larvae do not. Data on larval growth of neritimorph gastropods are needed to help determine which of these growth patterns for planktotrophic gastropod larvae is more derived. We cultured planktotrophic larvae from all three major gastropod clades with feeding larvae through delays of metamorphosis of 3–10 weeks. Larvae of the caenogastropod Euspira lewisii and the euthyneurans Haminoea vesicula (Opisthobranchia) and Siphonaria denticulata (Pulmonata) conformed to previously described growth patterns for their respective major clades. Furthermore, the caenogastropod continued to lengthen the prototroch (ciliary band for swimming and feeding) and to differentiate prospective post-metamorphic structures (gill filaments and radular teeth) during delayed metamorphosis. Larvae of the neritimorph Nerita atramentosa arrested shell growth during delayed metamorphosis but the radula continued to elongate, a pattern most similar to that of non-feeding larvae of Haliotis, a vetigastropod genus. Character mapping on a phylogenetic hypothesis suggests that large larval size and capacity for continued growth during delayed metamorphosis, as exhibited by some caenogastropods, is a derived innovation among feeding gastropod larvae. This novelty may have facilitated post-metamorphic evolution of predatory feeding using a long proboscis.     

Otolith microstructure and daily increment validation of marbled sole (<Emphasis Type="Italic">Pseudopleuronectes yokohamae</Emphasis>)

We examined the daily deposition of otolith increments of marbled sole (Pseudopleuronectes yokohamae) larvae and juveniles by rearing experiments, and estimated the growth pattern of wild larvae and juveniles in Hakodate Bay (Hokkaido Island, Japan). At 16°C, prominent checks (inner checks; ca. 19.8 µm in diameter) were observed on the centers of sagittae and lapilli extracted from 5-day-old larvae. On both otoliths, distinctive and regular increments were observed outside of the inner checks, and the slopes of regression lines between age and the number of increments (n_i) (for sagittae: n_i=0.98×Day–5.90; for lapillus: n_i=0.96×Day–5.70) did not significantly differ from 1. Inner check formations were delayed at lower temperature, and the inner checks formed 13 days after hatching at 8°C. Over 80% of larvae, just after their yolk-sac has been absorbed completely (stage C), had inner checks on both their otoliths. On the lapilli, other checks (outer check) formed at the beginning of eye migration (stage G). To validate the daily deposition of increments during the juvenile stage, wild captured P. yokohamae juveniles were immersed in alizarin complexone (ALC)-seawater solutions and reared in cages set in their natural habitat. After 6 days, the mean number of rings deposited after the ALC mark was 5.7. The age–body length relationship of wild P. yokohamae larvae and juveniles caught in Hakodate Bay was divided into three phases. In the larval period, the relationship was represented by a quadratic equation (notochord length=–0.010×Age²+0.682×Age–2.480, r²=0.82, P<0.001), and the estimated instantaneous growth was 0.38 mm day^–1 at 15 days, 0 mm day^–1 at 34 days and –0.12 mm day^–1 at 40 days. The age–body length relationship in the early juvenile stage (<50 days) and the late juvenile stage (>50 days) were represented by linear equations (standard length=0.055×Age+5.722 and standard length=0.345×Age–9.908, respectively). These results showed that the growth rates in the late larval periods and the early juvenile stage were lower than those in the early larval stage and late juvenile stage; during the slow growth period, energy appears to be directed towards metamorphosis rather than body growth. This study provided the information needed to use otolith microstructure analysis for wild marbled sole larvae and juveniles.Communicated by T. Ikeda, Hakodate     

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.     

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.     

Dissolved histamine: a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae

Many species of marine invertebrate larvae settle and metamorphose in response to chemicals produced by organisms associated with the adult habitat, and histamine is a cue for larvae of the sea urchin Holopneustes purpurascens. This study investigated the effect of histamine on larval metamorphosis of six sea urchin species. Histamine induced metamorphosis in larvae of three lecithotrophic species (H. purpurascens, Holopneustes inflatus and Heliocidaris erythrogramma) and in one planktotrophic species (Centrostephanus rodgersii). Direct comparisons of metamorphic rates of lecithotrophic and planktotrophic larvae in assays cannot be made due to different proportions of larvae being competent. Histamine (10 μM) induced metamorphosis in 95% of larvae of H. purpurascens and H. inflatus after 1 h, while the coralline alga Amphiroa anceps induced metamorphosis in 40–50% of these larvae. Histamine (10 μM) and A. anceps induced 40 and 80% metamorphosis, respectively, in the larvae of H. erythrogramma after 24 h. Histamine (10 μM) and the coralline alga Corallina sp. induced 30 and 70% metamorphosis, respectively, in the larvae of C. rodgersii after 24 h. No metamorphosis of any larval species occurred in seawater controls. Larvae of two planktotrophic species (Tripneustes gratilla and Heliocidaris tuberculata) did not metamorphose in response to histamine. Seagrasses, the host plants of H. inflatus, induced rapid metamorphosis in larvae of the two Holopneustes species, and several algae induced metamorphosis in C. rodgersii larvae. Histamine leaching from algae and seagrasses may act as a habitat marker and metamorphic cue for larvae of several ecologically important sea urchin species.     

Abundance of estuarine crab larvae is associated with tidal hydrologic variables

Abundances of brachyuran megalopae and juveniles were measured throughout consecutive tidal cycles during six 2 to 3 d sampling periods in summer 1992, and associated with rates of change of tidal hydrologic variables in the Newport River Estuary. Current speeds and rates of pressure change fitted sinusoidal (tidal) models well; however, rates of salinity and temperature change did not. Analysis of plankton samples taken during spring and neap tides showed peak abundances during nighttime rising tides for all taxonomic groups: Callinectes sapidus, Uca spp., Xanthidae, and Pinnixa spp. megalopae, and Pinnotheres spp. juveniles. Megalopal and juvenile abundances from time-intensive sampling were related to rates of changes in the hydrologic variables using stepwise logistic regression. No hydrologic variable accounted well for the presence of Uca spp. megalopae. Megalopal presence was best predicted by current speed for Pinnixa spp. megalopae, and rates of changes in pressure for xanthid megalopae and Pinnotheres spp. juveniles, and salinity for C. sapidus megalopae. These variables might act as cues causing megalopae to ascend into the water column at a particular point in the flooding tide, and subsequently descend to or near the bottom prior to ebb flow. In this way, larvae which develop on the continental shelf or lower estuary undergo transport up the estuary by behaviorally altering their swimming activity and depth concurrent with tidal changes.