Effects of egg size reduction and larval feeding on juvenile quality for a species with facultative-feeding development

In free-spawning marine invertebrates, larval development typically proceeds by one of two modes: planktotrophy (obligate larval feeding) from small eggs or lecithotrophy (obligate non-feeding) from relatively large eggs. In a rare third developmental mode, facultative planktotrophy, larvae can feed, but do not require particulate food to complete metamorphosis. Facultative planktotrophy is thought to be an intermediate condition that results from an evolutionary increase in energy content in the small eggs of a planktotrophic ancestor. We tested whether an experimental reduction in egg size is sufficient to restore obligate planktotrophy from facultative planktotrophy and whether the two sources of larval nutrition (feeding and energy in the egg) differentially influence larval survival and juvenile quality. We predicted, based on its large egg size, that a reduction in egg size in the echinoid echinoderm Clypeaster rosaceus would affect juvenile size but not time to metamorphosis. We reduced the effective size of whole (W) zygotes by separating blastomeres at the two- or four-cell stages to create half- (H) or quarter-size (Q) “zygotes” and reared larvae to metamorphosis, both with and without particulate food. Larvae metamorphosed at approximately the same time regardless of food or egg size treatment. In contrast, juveniles that developed from W zygotes were significantly larger, had higher organic content and had longer and more numerous spines than juveniles from H or Q zygotes. Larvae from W, H and Q zygotes were able to reach metamorphosis without feeding, suggesting that the evolution of facultative planktotrophy in C. rosaceus was accompanied by more than a simple increase in egg size. In addition, our results suggest that resources lost by halving egg size have a larger effect on larval survival and juvenile quality than those lost by withholding particulate food.     

The relationship between egg size and the duration of the facultative feeding period in marine invertebrate larvae

Feeding larvae of marine invertebrates fuel development from both endogenous egg energy and exogenous energy obtained from the planktonic environment. Although both sources of energy likely influence certain larval stages, only the effects of exogenous food have been well studied. Despite the lack of research on the effects of egg size on larval stages, investigators have hypothesized that egg size influences the duration of the facultative feeding stage—the stage in which larvae can feed but do not have to because development is still being fueled by egg energy. To test this hypothesis, we investigated six species of sand dollars with different sized eggs and quantified the duration of the larval facultative feeding period of each species by comparing when fed and starved larvae diverged in size. Regardless of whether phylogeny was taken into account, the duration of the facultative feeding period was positively correlated with egg size. We further determined that our conclusions were not sensitive to either our estimation of the duration of the facultative feeding period, or the branch lengths of the phylogeny we used. This relationship is likely a result of larger eggs being provisioned with more energy, and may affect how well larvae can cope with natural variability in food concentrations. Furthermore, our results support an assumption of a theoretical model developed to understand the evolution of different life-history strategies in marine invertebrate larvae, which suggests that this relationship has important evolutionary consequences.     

Egg size and the evolution of phenotypic plasticity in larvae of the echinoid genus Strongylocentrotus

Planktotrophic larvae grow by utilizing energy obtained from food gathered in the plankton. Morphological plasticity of feeding structures has been demonstrated in multiple phyla, in which food-limited larvae increase feeding structure size to increase feeding rates. However, before larvae can feed exogenously they depend largely on material contained within the egg to build larval structures and to fuel larval metabolism. Thus, the capacity for plasticity of feeding structures early in development may depend on egg size. Using the congeneric sea urchins Strongylocentrotus franciscanus and S. purpuratus, which differ in egg volume by 5-fold, I tested whether the degree of expression of feeding structure (larval arm length) plasticity is correlated with differences in the size of the egg. I experimentally manipulated egg size of S. franciscanus (the larger-egged species) by separating blastomeres at the 2-cell stage to produce half-sized larvae. I reared half-size and normal-size larvae under high and low food treatments for 20 days. I measured arm and body lengths at multiple ages during development and calculated the degree of plasticity expressed by larvae from all treatments. Control and unmanipulated S. franciscanus larvae (from ∼ 1.0 nl eggs) had significantly longer arms relative to body size and a significantly greater degree of plasticity than half-sized S. franciscanus larvae (from < 0.18 nl eggs), which in turn expressed a significantly greater degree of plasticity than S. purpuratus larvae (from ∼ 0.3 nl eggs). These results indicate that egg size affects larval arm length plasticity in the genus Strongylocentrotus; larger eggs produce more-plastic larvae both in an experimental and a comparative context. However, changes in egg size alone are not sufficient to account for evolved differences in the pattern of plasticity expressed by each species over time and may not be sufficient for the evolutionary transition from feeding to non-feeding.     

Reproductive strategies of marine benthic invertebrates revisited: facultative feeding by planktotrophic larvae

Fecundity-time models of reproductive strategies in marine invertebrates all predict that reproductive success is maximized only at the extreme levels of investment. Selection should drive egg sizes toward small eggs and planktotrophy or large eggs and lecithotrophy. The existence of two distinct larval types, feeding and nonfeeding, has been taken as confirmation of this prediction and has established the current paradigm for larval ecology. However, comparative and experimental evidence does not support the prediction that egg size is minimized in species with planktotrophic larvae. Recent discoveries have documented the existence of planktotrophs that have intermediate egg sizes, differing degrees of dependence on exogenous food, and differing capacities for facultative feeding. A fecundity-time model is presented that includes facultative larval feeding by dissociating the onset of feeding capability from the need for exogenous food. The facultative feeding model shows that reproductive success can be maximized at intermediate levels of investment per offspring between the minimum for development and the threshold for lecithotrophy, depending on the amount of food available to larvae and the intensity of planktonic mortality. A continuum of larval strategies is predicted.     

The effect of age at metamorphosis on the transition from larval to adult suspension‐feeding of the slipper limpet Crepidula fornicata

Slipper limpets use different ciliary feeding mechanisms as larvae and adults. Veliger larvae of Crepidula fornicata developed part of the adult feeding apparatus, including ctenidial filaments, neck lobe, and radula, before metamorphosis, but ctenidial feeding did not begin until well after loss of the larval feeding apparatus (velum) at metamorphosis. Earlier initiation of ctenidial feeding by individuals that were older larvae when metamorphosis occurred suggests continued development toward ctenidial feeding during delay of metamorphosis. Early juveniles produced a ciliary current through the mantle cavity and moved the radula in a grasping action before they began to capture algal cells on mucous strands or form a food cord. Either early juveniles could not yet form mucous strands or they delayed their production until development of other necessary structures. The neck canal for transporting food from ctenidium to mouth cannot develop before velar loss. In their first feeding, juveniles fed much like the adults except that the neck canal was less developed and the path of the food cord toward the mouth sometimes varied. As suspension feeders, calyptraeids lack the elaborations of foregut that complicate transition to juvenile feeding for many caenogastropods, but a path for the food cord must develop after velar loss. Why individuals can initiate ctenidial feeding sooner when they are older at metamorphosis is not yet known. The juveniles became sedentary soon after metamorphosis and were not observed to feed by scraping the substratum with the radula, in contrast to the first feeding by juveniles of another calyptraeid species, observed by Montiel et al. ( 2005 ).     

Effects of starvation on larval growth, survival, and metamorphosis of Manila clam Ruditapes philippinarum

The effects of starvation on larval growth, survival, and metamorphosis of Manila clam Ruditapes philippinarum at the temperature of 19.6–21.6 °C, the salinity of 34‰ and pH of 8.0 were investigated from May 18 to July 18, 2006. In this study, the early, middle and late umbo-veliger larvae with the shell lengths of 100, 140, and 190 μm were subject to temporary food deprivation for up to 4.5, 20, and 25d at 0.5, 4, 5d intervals, followed by refeeding for the remaining of a 24, 20, 25d period, respectively. The results suggested that the larvae should have shown considerable tolerance to starvation due to their endogenous and exterior nutrition material, for larvae and time to the point-of-no-return (PNR: the threshold point during starvation after which larvae could no longer metamorphose even if food is provided) were calculated to be 4.25, 17.54, and 22.17d. As the starvation period prolonged, the mean shell length of larvae starved got close to constants at 1.5, 4, and 15d after starvation, which were different for larvae at different stages when starvation began, survival of larvae decreased, and was lower in treatments starved earlier in development than those starved later, for the early, middle and late umbo-veliger larvae, after 4.5, 20 and 25d of starvation period, few larvaes were alive. After starvation period, the alive larvaes were able to metamorphose and had a capability of compensatory growth when refeeding was given. Starvation not only affected metamorphosis rate, but also caused the delay in the time to metamorphosis and the decrease in the metamorphosed sizes. For example, for the continuously-fed larvae, duration to metamorphosis was 20.7d, for larvae with a size of 100-μm starved for up to 4d, larvae with a size of 140-μm starved for up to 16d, larvae with a size of 190-μm starved for up to 20d, duration to metamorphosis were 29.7, 31.7, and 37.7d, the delay in duration to metamorphosis were 9, 11, and 17d, respectively. Furthermore, importance of nutrition material for maintaining larval survival during starvation and the compensatory growth on larvae at the same feeding time were discussed.     

Effects of food availability on larval development in the slipper limpet Crepidula onyx (Sowerby)

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C. onyx veligers. Larval development occurred only at 10⁴ cells ml⁻¹ and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10² cells ml⁻¹I. galbana or the blank control. At 10³ cells ml⁻¹, there was only a slight increase in shell length over 12 days. At 10⁴ cells ml⁻¹, about 40% of the larvae became competent in 18 days. At 10⁵ and 10⁶ cells ml⁻¹, more than 90% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis.     

Insights into How Longicorn Beetle Larvae Determine the Timing of Metamorphosis: Starvation-Induced Mechanism Revisited

Larvae of holometabolous insects must determine the timing of their metamorphosis. How they determine this timing has only been studied in detail for a few insect species. In a few species of Coleoptera, starvation is known to be a cue for metamorphosis, leading to the formation of smaller adults (starvation-induced pupation, SiP). We investigated the occurrence of SiP in the beetle Psacothea hilaris. When P. hilaris larvae were starved late in the feeding phase of the last (5th) instar, they exhibited typical SiP characterized by constancy of the time from food deprivation to pupation (TTP) irrespective of the body weight upon food deprivation or the length of prior feeding. In contrast, when larvae were starved early in the feeding phase, TTP decreased by roughly 1 day as the feeding became 1 day longer. The change in the response to starvation was estimated to occur on day 5.9 in the last instar. A series of refeeding experiments suggested that whereas SiP occurred readily in the larvae starved in the late feeding phase, activation of SiP was suspended until day 5.9 in the larvae starved early in the feeding phase. When P. hilaris larvae were fed continuously, they eventually ceased feeding spontaneously and pupated. The time length between spontaneous cessation of feeding and pupation was approximately equal to the TTP in SiP. This suggests that the same mechanism was activated by food deprivation in the late feeding phase and by spontaneous cessation of ad libitum feeding.     

Thyroid hormones determine developmental mode in sand dollars (Echinodermata: Echinoidea)

Evolutionary transitions in larval nutritional mode have occurred on numerous occasions independently in many marine invertebrate phyla. Although the evolutionary transition from feeding to nonfeeding development has received considerable attention through both experimental and theoretical studies, mechanisms underlying the change in life history remain poorly understood. Facultative feeding larvae (larvae that can feed but will complete metamorphosis without food) presumably represent an intermediate developmental mode between obligate feeding and nonfeeding. Here we show that an obligatorily feeding larva can be transformed into a facultative feeding larva when exposed to the thyroid hormone thyroxine. We report that larvae of the subtropical sand dollar Leodia sexiesperforata (Echinodermata: Echinoidea) completed metamorphosis without exogenous food when treated with thyroxine, whereas the starved controls (no thyroxine added) did not. Leodia sexiesperforata juveniles from the thyroxine treatment were viable after metamorphosis but were significantly smaller and contained less energy than sibling juveniles reared with exogenous food. In a second starvation experiment, using an L. sexiesperforata female whose eggs were substantially larger than in the first experiment (202+/-5 vs. 187+/-5 microm), a small percentage of starved L. sexiesperforata larvae completed metamorphosis in the absence of food. Still, thyroxine-treated larvae in this experiment completed metamorphosis faster and in much higher numbers than in the starved controls. Furthermore, starved larvae of the sand dollar Mellita tenuis, which developed from much smaller eggs (100+/-2 microm), did not complete metamorphosis either with or without excess thyroxine. Based on these data, and from recent experiments with other echinoids, we hypothesize that thyroxine plays a major role in echinoderm metamorphosis and the evolution of life history transitions in this group. We discuss our results in the context of current life history models for marine invertebrates, emphasizing the role of egg size, juvenile size, and endogenous hormone production for the evolution of nonfeeding larval development.     

The effects of nurse eggs and sibling interactions on the larval development of the poecilogonous annelid Boccardia proboscidea (Spionidae)

In poecilogony, different types of larvae are produced within the same species. Previous studies have suggested maternal control of the production of larval types; however, it is not clear which factors or mechanisms generate contrasting developmental patterns among siblings. The spionid polychaete Boccardia proboscidea produces within the same capsule adelphophagic larvae that eat nurse eggs and siblings and complete all or most of their development inside the capsule (Type A larvae), and larvae with little growth until they hatch as planktotrophic larvae (Type B larvae). In this study, we manipulated capsule content to explore the factors determining larval type in B. proboscidea and the role of extra‐embryonic maternal nutrition and sib–sib interaction in the developmental fate of offspring. When early larval stages were grown individually in vitro, with nurse eggs as the only food source, some of them remained small, while others continue developing into larger pre‐competent larvae by feeding on nurse eggs. This suggests that larval types in B. proboscidea are determined very early in development and are not solely the product of sib–sib interaction inside the capsule. However, our data also suggest that hatching size variability within larval types of a clutch depends on nurse egg availability. Type B larvae grew normally to metamorphosis when phytoplankton was available, but suffered high rates of cannibalism by Type A larvae. These results are consistent with the hypothesis that individual larval fates are determined very early in development and that once their fate is determined, hatching size and intracapsular survival are affected by maternal food provisioning and sibling interaction.     

Lipid and fatty acid composition of striped bass (Morone saxatilis) larvae during development

Changes in lipid class, fatty acid composition, protein, and dry and wet weights of fertilized eggs and developing larvae of striped bass (Morone saxatilis) fed with the live food, Artemia, were investigated. A decrease of wet and dry weights and moisture was observed at the beginning of the larval stage. Larvae regained the original moisture level, and wet and dry weights increased steadily after feeding. Total lipids decreased from 190 μg/egg in fertilized eggs to 151 μg/egg during hatching and increased after feeding. When total lipid contents were expressed as a percentage of larval dry weight, a decline of lipid did not occur until after feeding. Total protein, on the other hand, increased right after feeding, but there was some variation between days. Polar lipids increased significantly from 20 μg/egg at the egg stage to 199 μg/larva at 26 days post-hatching (DPH), 2 days before the onset of metamorphosis, while neutral lipids declined from 175 μg/egg to 80 μg/larva during the same time period. Wax/steryl esters decreased from 150 μg/egg in fertilized eggs to 32 μg/larva at 26 DPH. Triacylglycerols dropped from 21 μg/egg to 15 μg/larva before feeding and increased gradually after feeding. In contrast, the level of cholesterol increased 2–3-fold. There was a significant increase of phospholipids, particularly phosphatidylcholine in larvae after feeding. The fatty acid composition of fish larvae was significantly influenced by the diet, Artemia. There was an indication of catabolism of endogenous eicosapentaenoic and docosahexaenoic acids during metamorphosis.     

Changes in biomass and chemical composition during lecithotrophic larval development of the southern king crab, Lithodes santolla (Molina)

Changes in biomass and elemental composition (dry mass, W; carbon, C; nitrogen, N; hydrogen, H) were studied in the laboratory during complete larval and early juvenile development of the southern king crab, Lithodes santolla (Molina), formerly known as Lithodes antarcticus (Jacquinot). At 6±0.5 °C, total larval development from hatching to metamorphosis lasted about 10 weeks, comprising three demersal zoeal stages and a benthic megalopa, with mean stage durations of 4, 7, 11 and 47 days, respectively. No differences in development duration or mortality were observed in larvae either fed with Artemia sp. nauplii or unfed, indicating that all larval stages of L. santolla are lecithotrophic. First feeding and growth were consistently observed immediately after metamorphosis to the first juvenile crab stage. Regardless of the presence or absence of food, W, C, N and H decreased throughout larval development. Also the C:N mass ratio decreased significantly, from 7.7 at hatching to 4.1 at metamorphosis, indicating that a large initial lipid store remaining from the egg yolk was gradually utilized as an internal energy source, while proteins played a minor role as a metabolic substrate. In total, 56-58% of the initial quantities of C and H present at hatching, and 20% of N were lost during nonfeeding larval development to metamorphosis. Nine to ten percent of the initially present C, N and H were lost with larval exuviae, half of these losses occurring in the three zoeal stages combined and another half in the megalopa stage alone. Metabolic biomass degradation accounted for losses of about 47-50% in C and H but for only 10% in N. Hence, most of the losses in C and H reflected metabolic energy consumption (primarily lipid degradation), while about half of the losses in N and two thirds of those in W were due to larval exuviation. Complete independence from food throughout larval development is based on an enhanced maternal energy investment per offspring and on energy-saving mechanisms such as low larval locomotory activity and low exuvial losses. These traits are interpreted as bioenergetic adaptations to food-limited conditions in Subantarctic regions, where a pronounced seasonality of day length limits the period of primary production, while low temperatures enforce a long duration of pelagic development.     

Bioenergetics of abbreviated larval development in the bromeliad crab,Metopaulias depressus (Decapoda: Grapsidae)

• 1.1. Larvae of the bromeliad crab, Metopaulias depressus Rathbun, were reared in the laboratory, and changes in dry weight (W), ash-free W (AFW), carbon, nitrogen, hydrogen, protein, lipid, carbohydrates and respiration rate were measured during development from hatching to metamorphosis.
• 2.2. Development was successful in rain-water from bromeliads (pH < 5–6), but not in river water from the same region (pH 8). It is abbreviated, with two non-feeding zoeal stages (2.5–3.5 days each) and a feeding megalopa (8.5–10 days). Development to metamorphosis can also be completed in the absence of food (facultative lecithotrophy).
• 3.3. Dry weight and other absolute biomass values per individual vary significantly between different hatches, whereas changes in the relative (% of W or AFW) composition follow quite invariable patterns: ash increases from hatching through the first part of megalopa development, organic biomass decreases concurrently.
• 4.4. Elemental and biochemical data show that lecithotrophy of the zoeal stages as well as continued endotrophic development in the megalopa depend chiefly on degradation of lipid reserves and less on protein. No significant growth was observed in organic constituents when food was available, but without food the megalopa reached metamorphosis with only half the lipid and less than two thirds the protein of fed siblings.
• 5.5. The relationship between C and lipid is similar in M. depressus larvae as in planktotrophic marine crab larvae, whereas that between N and protein differs; it indicates the presence of unusually large quantities of unidentified non-protein N.
• 6.6. Exuvial losses of late premoult biomass or energy are very low in the zoeal stages (2 and 3%), but increase in the megalopa (16% in W, 10% in C, 7–8% in N, H and energy).
• 7.7. Respiration rate per individual increases gradually during larval development (0.6–0.8 μg O₂/hr). Starved megalopa larvae reveal lower individual but higher W-specific metabolism than fed larvae.
• 8.8. Bioenergetic traits of abbreviated larval development are discussed in relation to those known from regular (planktotrophic marine) development of brachyuran crabs. M. depressus is highly adapted to life and development in a physically extreme terrestrial environment.

     

Facultative Second Oviposition as an Adaptation to Egg Loss in a Semelparous Crab Spider

Semelparity is prevalent in arthropod species that exhibit maternal care. Previous hypotheses postulated that long‐term maternal care constrains future reproduction in females, leading to the evolution of semelparity. Nevertheless, females may occasionally lose all or part of their offspring because of predation or other causes. Where females lose the first egg mass for any reason, the potential for females to produce an additional egg mass could be adaptive. This potential may be found widely among semelparous arthropods as a conditional strategy. We tested this hypothesis using the crab spider Lysiteles coronatus whose females guard their egg mass against predators. L. coronatus females did not consume food during the 40‐d guarding period; this resulted in a 30.2% loss in their weight. When the females were separated from their eggs immediately after oviposition and were provided with food, they resumed feeding and their ovaries redeveloped. Dissection of guarding females indicated that their ovaries developed temporarily during egg guarding and that the developed ovaries were subsequently reabsorbed. These results suggest that the females maintain the potential to produce a second egg mass in case of egg loss, but that this potential declines towards the end of the guarding period. Field observations showed that a small fraction of the females oviposited in late July, when most females had completed egg guarding. The size of the late broods was similar to the oocyte numbers that we found in the females fed in the laboratory. This result suggests that a few females produced a second egg mass after they had lost the first one. Thus, we suggest that facultative second oviposition in L. coronatus females has evolved as an adaptation to egg loss, and that the development of ovaries during the guarding period is intrinsically programmed for compensatory oviposition.     

Effect of changes in resource level on age and size at metamorphosis in Hyla squirella

Recent experiments suggest that timing of metamorphosis is fixed during development in some anurans, insects, and freshwater invertebrates. Yet, these experiments do not exclude a growth rate optimization model for the timing of metamorphosis. I manipulated food resources available to larvae of squirrel treefrogs (Hyla squirella) to determine if there is a loss of plasticity in duration of larval period during development and to critically test growth rate models for the timing of metamorphosis. Size-specific resource levels for individual tadpoles were switched from low to high or high to low at three developmental stages spaced throughout larval development. The effects of changes in resource availability on larval period and mass at metamorphosis were measured. Switching food levels after late limb bud development did not significantly affect larval period in comparison to constant food level treatments. Therefore, developmental rate in H. squirella is better described by a fixed developmental rate model, rather than a growth rate optimization model. The timing of fixation of developmental rate in H. squirella is similar to that found in other anuran species, suggesting a taxonomically widespread developmental constraint on the plasticity of larval period duration. Mass at metamorphosis was not significantly affected by the timing of changes in food levels; the amount of food available later in development determined the size at metamorphosis. Larval period and mass at metamorphosis were negatively correlated in only one of two experiments, which contrasts with the common assumption of a phenotypic trade-off between decreased larval period and increased mass at metamorphosis. Received: 19 August 1996 / Accepted: 20 June 1997     

Effects of GABA and epinephrine on the settlement and metamorphosis of the larvae of four species of bivalve molluscs

Competent larvae of different marine bivalve species were treated with GABA and epinephrine at different concentrations and times of exposure to test the ability of the drugs to induce settlement and metamorphosis. GABA induced both settlement and metamorphosis in the mussel Mytilus galloprovincialis, the clams Venerupis pullastra and Ruditapes philippinarum and the oyster Ostrea edulis. Maximum induction of settlement (>39%) was achieved after exposure of V. pullastra larvae to 10⁻⁴ M GABA; this concentration of GABA also induced the highest percentages of metamorphosis in the four species studied. Epinephrine was identified as an active inducer of settlement and metamorphosis in bivalve molluscs. Exposure to 10⁻⁵ M epinephrine induced significant levels of settlement in Mytilus, Venerupis and Ostrea. In contrast, epinephrine failed to induce settlement behaviour in Ruditapes. Maximum induction of metamorphosis was produced by 10⁻⁵ M epinephrine in mussels, clams and oysters; Ruditapes showed the highest percentage of metamorphosis (>78%). This is the first report in which the involvement of GABA in the settlement and metamorphosis of bivalve molluscan larvae is demonstrated. It was also recognised that epinephrine plays a role not only in inducing metamorphosis but also in initiating settlement.     

The relationship among egg size,density and food level on larval development in the wood frog (Rana sylvatica)

Summary Although inter- and intraspecific variation in egg size among amphibians has been well documented, the relationship between egg size and fitness remains unclear. Recent attempts to correlate egg size intraspecifically with larval developmental patterns have been equivocal. In this study the development of larvae derived from large eggs and small eggs, from a single population in Maryland were compared under a range of food levels and larval population densities. Both food level and density had significant effects on the length of the larval period and size at metamorphosis. However, the response among larvae derived from different egg sizes was not additive. At low densities and high food levels, larvae from small eggs had longer larval periods and a larger size at metamorphosis than larvae derived from large eggs. In contrast, at high densities larvae from small eggs had longer developmental periods but were smaller at metamorphosis than larvae from large eggs. In addition, larvae from small eggs were more sensitive to density irrespective of food level. These results suggest that optimal egg size is correlated with environmental factors, which may explain the maintenance of both geographic and within population variation in egg size commonly observed in amphibians.     

Observations on development,larval growth and metamorphosis of four species of aplysiidae (gastropoda: Opisthobranchia) in laboratory culture

The development of simple, reliable techniques for the laboratory culture of aplysiid gastropods through their complete life cycle, has enabled us to study the larval biology, metamorphosis, and early juvenile development of these animals. Egg masses, duration of the embryonic phase, veligers, and larval growth and development are described for four species of Hawaiian Aplysiidae, namely, Aplysia dactylomela Rang, Aplysia Juliana Quoy and Gaimard, Dolabella auricularia (Lightfoot) and Stylocheilus longicauda (Quoy and Gaimard). Metamorphosis and early juvenile development of A. Juliana are described in detail with additional comments on these processes in the other three species. Length of the embryonic phase and size of the veliger at hatching are a function of the size of the uncleaved egg. All four species develop planktotrophically and have ≈ 30-day larval phases. In each species the larval phase includes a period of rapid shell growth to a species-specific size followed by a non-growth period during which other morphological developments occur to culminate in metamorphic competence. The larvae of each species metamorphose preferentially on a particular species of benthic algae. The events of metamorphosis require 2 to 4 days for completion and transform the planktonic filter-feeding larva into a benthic, radular-feeding juvenile. Postlarval development includes growth of the shell, parapodia, oral tentacles, rhinophores, anal siphon, and structures of the mantle cavity.     

EFFECTS OF EGG SIZE ON POSTLARVAL PERFORMANCE: EXPERIMENTAL EVIDENCE FROM A SEA URCHIN

Many life-history and developmental studies of marine invertebrates assume that eggs of species with nonfeeding larvae are large because they provide materials for rapid development. Using the sea urchin Heliocidaris erythrogramma which has 400 μm eggs and nonfeeding larvae, we removed an acellular, lipid-rich component from the blastula equivalent to ca. 40% of the egg volume and ca. 50% of the organic mass. Experimentally manipulated, reduced-lipid larvae survived well, developed in the usual time (3.5 d), and high percentages of the original numbers metamorphosed into anatomically normal juveniles. Control juveniles were larger at metamorphosis, grew more, and survived longer than siblings that lacked this lipid-rich material. Moderate increases in egg size in species with nonfeeding larvae may enhance postlarval performance significantly and therefore may reflect selection on early juvenile traits. The contrasts of our results and comparable experiments with feeding larvae suggests that egg size may play fundamentally different roles in species with feeding and nonfeeding larvae. The accommodation of materials reserved for the juvenile stage should be considered among hypotheses on evolutionary modification of developmental patterns.     

How do changes in parental investment influence development in echinoid echinoderms?

SUMMARY Understanding the relationship between egg size, development time, and juvenile size is critical to explaining patterns of life-history evolution in marine invertebrates. Currently there is conflicting information about the effects of changes in egg size on the life histories of echinoid echinoderms. We sought to resolve this conflict by manipulating egg size and food level during the development of two planktotrophic echinoid echinoderms: the green sea urchin, Strongylocentrotus droebachiensis and the sand dollar, Echinarachnius parma . Based on comparative datasets, we predicted that decreasing food availability and egg size would increase development time and reduce juvenile size. To test our prediction, blastomere separations were performed in both species at the two-cell stage to reduce egg volume by 50%, producing whole- and half-size larvae that were reared to metamorphosis under high or low food levels. Upon settlement, age at metamorphosis, juvenile size, spine number, and spine length were measured. As predicted, reducing egg size and food availability significantly increased age at metamorphosis and reduced juvenile quality. Along with previous egg size manipulations in other echinoids, this study suggests that the relationship between egg size, development time, and juvenile size is strongly dependent upon the initial size of the egg.