Showing papers in "The Biological Bulletin in 2009"

Pelagic Larval Duration and Dispersal Distance Revisited

Abstract: I present dispersal distances for 44 species with data on propagule duration (PD) for 40 of these. Data were combined with those in Shanks et al. (2003; Ecol. Appl. 13: S159 -S169), providing information on 67 species. PD and dispersal distance are correlated, but with many exceptions. The distribution of dispersal distances was bimodal. Many species with PDs longer than 1 day dispersed less than 1 km, while others dispersed tens to hundreds of kilometers. Or- ganisms with short dispersal distances were pelagic briefly or remained close to the bottom while pelagic. Null models of passively dispersing propagules adequately predict dis- persal distance for organisms with short PDs (1 day), but overestimate dispersal distances for those with longer PDs. These models predict that propagules are transported tens of kilometers offshore; however, many types remain within the coastal boundary layer where currents are slower and more variable, leading to lower than predicted dispersal. At short PDs, dispersal distances estimated from genetic data are similar to observed. At long PDs, genetic data generally overestimate dispersal distance. This discrepancy is proba- bly due to the effect of rare individuals that disperse long distances, thus smoothing genetic differences between pop- ulations. Larval behavior and species' life-history traits can play a critical role in determining dispersal distance.

Quantifying the “Bio-” Components in Biophysical Models of Larval Transport in Marine Benthic Invertebrates: Advances and Pitfalls

Abstract: Biophysical models are being used increasingly, both as predictive tools of larval dispersal for a particular system and for general evaluation of the role of different factors in larval transport. In the results of such models, larval duration, mortality, and behavior in the water column have exhibited pronounced effects on larval dispersal of marine benthic invertebrates. The parameterization of these processes has broadly reflected values from laboratory experiments, but the accuracy of these values is unknown. The pelagic larval duration used in models should be determined by laboratory, or preferably field, studies and should incorporate environmentally dependent variability. For mortality, in situ estimates are now feasible and, likely, more accurate than the currently used values. Larval behavior can be measured in the field, by high-frequency sampling of distributional changes relative to features in the water column or by controlled larval releases in tractable systems. To successfully validate the outcomes of these models, we must either improve our techniques for measuring larval abundance at the end of larval transport immediately before settlement, or incorporate components for settlement into the models. We must also address the mismatch in sampling resolution between biological and physical processes. If used with caution, this powerful approach can significantly advance our understanding of larval transport.

Egg size as a life history character of marine invertebrates: Is it all it's cracked up to be?

Abstract: Egg size is one of the most important aspects of the life history of free-spawning marine organisms, and it is correlated with larval developmental mode and many other life-history characters. Egg size is simple to measure and data are available for a wide range of taxa, but we have a limited understanding of how large and small eggs differ in composition; size is not always the best measure of the characters under selection. Large eggs are generally consid- ered to reflect increased maternal investment, but egg size alone can be a poor predictor of energetic content within and among taxa. We review techniques that have been used to measure the energetic content and biochemical makeup of invertebrate eggs and point out the strengths and diffi- culties associated with each. We also suggest a number of comparative and descriptive approaches to biochemical constituent analysis that would strengthen our understand- ing of how natural selection shapes oogenic strategies. Fi- nally, we highlight recent empirical research on the intrinsic factors that drive intraspecific variation in egg size. We also highlight the relative paucity of these data in the literature and provide some suggestions for future research directions.

Linking Thermal Tolerances and Biogeography: Mytilus edulis (L.) at its Southern Limit on the East Coast of the United States

Abstract: Temperature is a major factor contributing to the latitudinal distribution of species. In the Northern Hemisphere, a species is likely to be living very close to its upper thermal tolerance limits at the southern limit of its biogeographic range. With global warming, this southern limit is expected to shift poleward. Moreover, intertidal ecosystems are expected to be especially strongly affected, mostly due to their large daily and seasonal variations in temperature and exposure. Hence, these are model systems in which to conduct experiments examining the ecological effects of climate change. In this study we determined the upper lethal thermal limits, for both air and water, of the blue mussel Mytilus edulis via laboratory experiments. Tolerances vary seasonally, with a difference between media of 0.7 °C in June and 4.8 °C in November, as well as a decrease with multiple exposures. Measured lethal limits were then compared to field measurements of environmental temperature and concurrent measurements of ...

Elevated Temperature Affects Development, Survivorship, and Settlement of the Elkhorn Coral, Acropora palmata (Lamarck 1816)

Abstract: Elevated seawater temperatures during the late summer have the potential to negatively affect the development and survivorship of the larvae of reef corals that are reproductive during that time of year. Acropora palmata, a major Caribbean hermatype, reproduces annually during August and September. A. palmata populations have severely declined over the past three decades, and recovery will require high recruitment rates. Such recruitment will be limited if larval supply is reduced by elevated temperatures. The effects of elevated temperatures on development, survival, and larval settlement of A. palmata were investigated by culturing newly fertilized eggs at temperatures ranging from 27.5 to 31.5 degrees C. Development was accelerated and the percentage of developmental abnormalities increased at higher temperatures. Embryo mortality peaked during gastrulation, indicating that this complex developmental process is particularly sensitive to elevated temperatures. Larvae cultured at 30 and 31.5 degrees C experienced as much as an 8-fold decrease in survivorship compared to those at 28 degrees C. Additionally, settlement was 62% at 28 degrees C compared to 37% at 31.5 degrees C. These results indicate that embryos and larvae of A. palmata will be negatively affected as sea surface temperatures continue to warm, likely reducing recruitment and the recovery potential of A. palmata on Caribbean reefs.

Shaping the Things to Come: Ontogeny of Lophotrochozoan Neuromuscular Systems and the Tetraneuralia Concept

Abstract: Despite the large variation in adult bodyplan phenotypes, a worm-shaped morphology is considered plesiomorphic for both Lophotrochozoa and Bilateria. Although almost all larval and adult lophotrochozoan worms have serially arranged ring muscles in their body wall, a comparison of their ontogeny reveals no less than six different developmental pathways that lead to this homogenous arrangement of ring muscles. However, in all taxa, with the exception of chaetodermomorph molluscs and the segmented annelids, ring muscle development starts with synchronous formation of certain pioneer myocytes, which is thus considered basal for Lophotrochozoa. Recent studies on spiralian neurogenesis revealed remnants of ancestral segmentation in echiurans and sipunculans, thus confirming molecular phylogenetic studies that propose a close relationship of these three taxa. Larval entoprocts exhibit a mosaic of larval and adult molluscan characters and, among other apomorphies, share with polyplacophoran Mollusca a complex larval apical organ and a tetraneurous nervous system, strongly suggesting a monophyletic assemblage of Entoprocta and Mollusca. The term Tetraneuralia is proposed herein for this lophotrochozoan clade. Overall, formation of the lophotrochozoan neuromuscular bodyplan appears as a highly dynamic process on both the ontogenetic and the evolutionary timescales, highlighting the importance of insights into these processes for reconstructing ancestral bodyplan features and phylogenetic relationships.

Spawning, Development, and the Duration of Larval Life in a Deep-Sea Cold-Seep Mussel

Abstract: We describe culturing techniques and development for the cold-seep mussel "Bathymodiolus" childressi, the only deep-sea bivalve for which development has been detailed. Spawning was induced in mature mussels by injection of 2 mmol l(-1) serotonin into the anterior adductor muscle. The mean egg diameter is 69.15 +/- 2.36 microm (+/-S.D.; n = 50) and eggs are negatively buoyant. Cleavages are spiral and at 7-8 degrees C occur at a rate of one per 3-9 h through hatching, with free-swimming blastulae hatching by 40 h and shells beginning to develop by day 12. When temperature was raised to 12-14 degrees C after hatching, larvae developed to D-shell veligers by day 8 without being fed. Egg size and larval shell morphology indicate that "B." childressi has a planktotrophic larva, but we did not observe feeding in culture. Wide distribution of this species throughout the Gulf of Mexico and amphi-Atlantic distributions of closely related congeners suggest that larvae may spend extended periods in the plankton. Duration of larval life was estimated for "B." childressi by comparing calculated settlement times to known spawning seasons. These estimates suggest variability in the larval duration, with individuals spending more than a year in the plankton.

Lectins associated with the feeding organs of the oyster Crassostrea virginica can mediate particle selection.

Abstract: Despite advances in the study of particle selection in suspension-feeding bivalves, the mechanisms upon which bivalves rely to discriminate among particles have not been elucidated. We hypothesized that particle sorting in suspen- sion-feeding bivalves could be based, in part, on a biochemical recognition mechanism mediated by lectins within the mucus that covers the feeding organs. Using Crassostrea virginica, the Eastern oyster, our investigations demonstrated that lectins from oyster mucus can specifically bind several microalgal species as well as different types of red blood cells (RBC), triggering their agglutination. Agglutination of microalgal spe- cies and RBC varied with the source of mucus (gills vs. labial palps). Hemagglutination and hemagglutination inhibition as- says emphasized that mucus contains several lectins. In feed- ing experiments, Nitzschia closterium and Tetraselmis macu- lata were separately incubated with mucus before being fed to oysters. Results showed that pre-treating these microalgae with mucus significantly alters the ability of oysters to sort particles. In another experiment, oysters were fed a mixture of micro- spheres coated with either bovine serum albumin (BSA) or glucosamide-BSA. Results show that oysters preferentially ingest microspheres with bound carbohydrates, highlighting probable interactions between lectins and carbohydrates in the mechanisms of microalgae recognition. This study confirms the presence of lectins in mucus that covers the feeding organs of oysters and suggests a new concept with regard to particle processing by suspension-feeding bivalves: specific interac- tions between carbohydrates on the surface of particles and

Not my "type": larval dispersal dimorphisms and bet-hedging in opisthobranch life histories.

Abstract: When conditions fluctuate unpredictably, selec- tion may favor bet-hedging strategies that vary offspring characteristics to avoid reproductive wipe-outs in bad sea- sons. For many marine gastropods, the dispersal potential of offspring reflects both maternal effects (egg size, egg mass properties) and larval traits (development rate, habitat choice). I present data for eight sea slugs in the genus Elysia (Opisthobranchia: Sacoglossa), highlighting potentially adaptive variation in traits like offspring size, timing of metamorphosis, hatching behavior, and settlement response. Elysia zuleicae produced both planktotrophic and lecitho- trophic larvae, a true case of poecilogony. Both intracapsu- lar and post-hatching metamorphosis occurred among clutches of "Boselia" marcusi, E. cornigera, and E. crispata, a dispersal dimorphism often misinterpreted as poecilogony. Egg masses of E. tuca hatched for up to 16 days but larvae settled only on the adult host alga Halimeda, whereas most larvae of E. papillosa spontaneously meta- morphosed 5-7 days after hatching. Investment in extra- capsular yolk may allow mothers to increase larval size relative to egg size and vary offspring size within and among clutches. Flexible strategies of larval dispersal and offspring provisioning in Elysia spp. may represent adapta- tions to the patchy habitat of these specialized herbivores, highlighting the evolutionary importance of variation in a range of life-history traits.

Gill Development and Its Functional and Evolutionary Implications in the Blue Mussel Mytilus edulis (Bivalvia: Mytilidae)

Abstract: Study of gill development in bivalve larvae and postlarvae provides information on the evolution of this organ and feeding mechanisms of early stages. Scanning electron microscopy was used to document the development of the filibranch homorhabdic gill in hatchery-reared larval, postlarval, and juvenile Mytilus edulis. Four key stages were identified during gill development: (1) transfer of the particle collection function from velum to gill at metamorphosis, with subsequent elongation of the gill filaments to form a gill basket, with complete frontal ciliation; (2) reflection of the inner demibranchs, and transition to a V-shaped gill; (3) delayed development of the outer demibranchs, occuring simultaneously along the gill axis, with transition to the adult final W-shape; and (4) formation of the ventral particle grooves and concomitant acquisition of dense abfrontal ciliation. These key stages signal shifts in the mechanisms of particle processing during the early development of M. edulis. Gill development in the homorhabdic filibranch M. edulis was similar to that of the early homorhabdic stages of the heterorhabdic filibranchs studied to date (Pectinidae), but different from that of the pseudolamellibranchs (Ostreidae), suggesting divergent evolution of this character. Similarly, the systems responsible for gill cohesion and structural integrity are common to both the homorhabdic and heterorhabdic filibranchs, suggesting evolutionary proximity, but they are patently different from those of the eulamellibranchs and pseudolamellibranchs, suggesting evolutionary divergence.

Molluscan Larvae: Pelagic Juveniles or Slowly Metamorphosing Larvae?

Abstract: Asking the right questions about evolution of development, larval morphology, and life history requires knowledge of ancestral state. Two hypotheses dominate current opinion about the ancestral life cycle of bilaterians: the "larva-first" and the "intercalation" hypotheses. Until recently, the larva-first hypothesis was preeminent. This proposes that the original indirect life cycle of bilaterians included a planktotrophic larva followed by a benthic adult. Phylogenetic evidence suggests that a planktotrophic larva is plesiomorphic for echinoderms. A preponderance of de- velopmental studies on echinoderms may have fostered a tendency to extrapolate conclusions about echinoderm de- velopment to other clades, particularly the concept that larval and juvenile/adult bodies are mostly separate entities. However, some of the recent reconstructions of bilaterian phylogeny suggest that nonfeeding larvae may have been ancestral for bilaterians, and these may have been interca- lated into a life cycle that was originally direct. I review comparative data on molluscan development that suggests the trochophore-like stage is little more than a gastrula with transient structures (prototroch and apical sensory organ) to allow a temporary planktonic phase during development. Most lineage founder cells of molluscan embryos generate progeny that develop through the veliger stage into struc- tures of the juvenile, which becomes benthic when the prototroch and apical sensory organ are lost. In light of this, the model of separate larval and juvenile bodies with the latter developing from nests of multipotent cells within the larva is inappropriate for molluscs. The intercalation hy- pothesis may be a better model for interpreting development of molluscs and other lophotrochozoans.

Red Fluorescent Protein Responsible for Pigmentation in Trematode-Infected Porites compressa Tissues

Abstract: . Reports of coral disease have increased dramatically over the last decade; however, the biological mechanisms that corals utilize to limit infection and resist disease remain poorly under...

Larval biology of the crab Rhithropanopeus harrisii (Gould): a synthesis.

Abstract: This synthesis reviews the physiological ecol- ogy and behavior of larvae of the benthic crab Rhithropan- opeus harrisii, which occurs in low-salinity areas of estu- aries. Larvae are released rhythmically around the time of high tide in tidal estuaries and in the 2-h interval after sunset in nontidal estuaries. As in most subtidal crustaceans, the timing of larval release is controlled by the developing embryos, which release peptide pheromones that stimulate larval release behavior by the female to synchronize the time of egg hatching. Larvae pass through four zoeal stages and a postlarval or megalopal stage that are planktonic before metamorphosis. They are retained near the adult population by means of an endogenous tidal rhythm in vertical migration. Larvae have several safeguards against predation: they undergo nocturnal diel vertical migration (DVM) and have a shadow response to avoid encountering predators, and they bear long spines as a deterrent. Photo- responses during DVM and the shadow response are en- hanced by exposure to chemical cues from the mucus of predator fishes and ctenophores. The primary visual pig- ment has a spectral sensitivity maximum at about 500 nm, which is typical for zooplankton and matches the ambient spectrum at twilight. Larvae can detect vertical gradients in temperature, salinity, and hydrostatic pressure, which are used for depth regulation and avoidance of adverse envi- ronmental conditions. Characteristics that are related to the larval habitat and are common to other crab larval species are considered.

Behaving in the dark: locomotor, chromatic, postural, and bioluminescent behaviors of the deep-sea squid Octopoteuthis deletron young 1972.

Abstract: Visual behaviors are prominent components of intra- and interspecific communication in shallow-water cephalopods. Meso- and bathypelagic cephalopods were believed to have limited visual communication, other than bioluminescence, due to the reduced illumination at depth. To explore potential visual behaviors in mesopelagic squid, we used undersea vehicles to observe 76 individuals of Octopoteuthis deletron. In contrast to predictions, we found this species capable of a variety of visually linked behaviors not previously reported for a deep-ocean cephalopod. The resultant ethogram describes numerous chromatic, postural, locomotor, and bioluminescent behavioral components. A few common body patterns-the whole appearance of the individual involving multiple components-are characterized. The behaviors observed from individual squid were compared using a Non-metric Multi-Dimensional Scaling (NMDS) ordination, onto which hydrographic and observation parameters were mapped. Observation length, specimen collection, and contact with the vehicle affected which behaviors were performed. A separate NMDS, analyzing the body patterns, indicated that these sets of behavioral components could be visualized as groups within the NMDS ordination. While the functional roles of the behaviors described are not yet known, our findings of numerous behaviors in O. deletron clearly indicate that bioluminescence is not the sole method of visual communication by deep-sea squid.

Transcriptome analysis of the circadian regulatory network in the coral Acropora millepora.

Abstract: Entrained circadian rhythms control many biological processes. These cyclical systems are synchronized by environmental signals but continue to free run for a considerable time when the synchronizing stimuli are removed. In scleractinian corals that reproduce by broadcast spawning, timing processes are essential in ensuring successful fertilization. It is not known whether spawn timing is regulated directly by environmental signals or if it is entrained and regulated by circadian or circalunar rhythms. The genetic components of circadian systems have been studied in considerable detail in microbes, plants, and animals. To identify potential participants in regulating scleractinian circadian systems, we have undertaken a bioinformatic analysis of the transcriptome of larvae of the coral Acropora millepora. Acropora contigs assembled from more than 600,000 sequencing reads from larval mRNA were searched via NCBI BLAST for matches to 24 key insect and mammalian circadian genes. Matches were found to all quer...

Mass culture and characterization of tumor cells from a naturally occurring invertebrate cancer model: applications for human and animal disease and environmental health.

Abstract: On the northeastern coast of the United States and Canada, Mya arenaria, the soft shell clam, develops a diffuse, hemopoetic tumor (a fatal leukemia-like cancer) resulting from inactivation of p53-like family member pro- teins.These malignant cells provide a model for an unrelated set of human cancer cells that are also characterized by mortalin-based cytoplasmic sequestration of wild-type p53 tumor suppressor protein (mortalin is the mitochondrial Hsp70 protein). Here we describe methods for mass culture and long-term storage of tumor cells from this cancer. These are the first successful efforts at maintaining malignant cells from any marine invertebrate in vitro. Following passage (subculture), these cultures undergo transition from primary cultures to non-immortalized cell lines that continue to proliferate and do not re-differentiate the normal hemocyte phenotype. We also characterize normal clam hemocytes and the pathology of cancerous clam hemocytes in vitro and in vivo using light and electron microscopy, cyto- and immunocytochemistry, molecular biology, and a phagocy- tosis assay. Our protocols provide biomedical and environ- mental researchers with ready access to this naturally oc- curring cancer model. We discuss the clam cancer model regarding (a) human health and disease; (b) animal health, disease, and aquaculture; (c) environmental health monitor- ing; and (d) future research directions.

Gonad Development During the Early Life of Octopus maya (Mollusca: Cephalopoda)

Abstract: Gonad development during the early life of Octopus maya is described in terms of histological, mor- phometric, oocytes growth, and somatic-oocyte relationship data obtained from octopus cultured at the UMDI-UNAM, in Sisal, Yucatan, Mexico. This study is the first publication on gonad development during the early life of Octopus maya. A total of 83 O. maya specimens were used; their sizes ranged from 6.5 to 76 mm of total length (TL), 4 to 28 mm of dorsal mantle length (DML), 2.5 to 20 mm of ventral mantle length (VML), and 0.0180 to 7.2940 g of fixed body weight (fBW). Animals were weighed and measured only after preservation. A loss of 10% of living weight was estimated for juvenile octopuses after formalin preservation. The relation of length to weight (VML, DML, TL/fBW) pooled for both sexes had a strong positive correlation (r), as shown by a potential power function that was quite close to 1. Compound images were produced from numerous microscopic fields. The histological examination revealed that, 4 months after hatching, male octopus (24.5 mm DML and 7.2940 g fBW) were in gonad stages 2 (maturing) to 3 (mature), with spermatogonia and spermatocytes in the tu- bule wall and abundant spermatids and spermatozoa in the central lumen of the seminiferous tubules, suggesting the occurrence of different phases of gonad development at different maturity stages. In contrast, females (22.5 mm DML and 4.8210 g fBW) at the same time since hatching were immature (stage 1), with many oogonia, few oocytes, and germinal epithelium. This suggests that males reach maturity earlier than females, indicating a probable onset of maturity for males at around 4 months of culture o r8go f wet body weight. Our results indicate the possibility that the size-at-weight can be recognized early with a degree of certainty that allows the sexes to be separated for culture purposes; but more detailed studies on reproduction in re- lation to endocrinology and nutrition are needed.

Impacts of suspended sediments on fertilization, embryonic development, and early larval life stages of the pacific herring, Clupea pallasi.

Abstract: Pacific herring reproduce in the San Francisco Bay estuary during times of the year when suspended sed- iment loads are highest due to freshwater input, yet little is known about the effects of sediment on herring early life stages. During the first 2 h after eggs contacted water, embryos were adhesive and susceptible to having sediment particles attach permanently to the chorion. Treatment with suspended San Francisco Bay dredged sediments at ecolog- ically relevant concentrations of 250 or 500 mg/l during this time period increased self-aggregation of the eggs and led to sublethal and lethal effects. After the firs t2hi nwater, sediments that contacted embryos did not attach to chorions and did not have an observable impact. Sediment treatment during the first 2 h was not linked statistically to declines in fertilization or total larval hatch rate, but it did produce significant sublethal effects that included increases in pre- cocious larval hatch and higher percentages of abnormal larvae, as well as an increase in larval mortality.

Analysis of the Intermediary Metabolism of a Reductive Chemoautotroph

Abstract: All extant life forms depend, directly or indirectly, on the autotrophic fixation of the dominant elements of the biosphere: carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur. We have earlier presented the canonical network of reactions that constitute the anabolism of a reductive chemoautotroph. Separating this network into subgraphs reveals several empirical generalizations: (1) acetate (acetyl-CoA), pyruvate, phosphoenol pyruvate, oxaloacetate, and 2-oxoglutarate serve as universal starting points for all pathways leading to the universal building blocks-20 amino acids and 4 ribonucleotide triphosphates; (2) all pathways are anabolic; (3) all reactions operate by complete utilization of outputs with no molecules left behind as waste, ensuring conservation of information; (4) the core metabolome of 120 compounds is acidic, consisting of compounds containing phosphoric or carboxylic acid or both; and (5) the core network is both brittle-vulnerable to a single break-and robust-having persisted for 4 billion years. Preliminary analysis of the chemical reactions and resultant structures reveals (a) a sparseness among possible molecular structures; (b) subdomains in the network; and (c) restriction of anabolism to a small set of rudimentary organic reactions with limited diversity in chemical mechanisms. These generalizations have implications for biogenesis and trophic ecology.

Functional morphology and fluid interactions during early development of the scyphomedusa Aurelia aurita.

Abstract: Scyphomedusae undergo a predictable ontogenetic transition from a conserved, universal larval form to a diverse array of adult morphologies. This transition entails a change in bell morphology from a highly discontinuous ephyral form, with deep clefts separating eight discrete lappets, to a continuous solid umbrella-like adult form. We used a combination of kinematic, modeling, and flow visualization techniques to examine the function of the medusan bell throughout the developmental changes of the scyphomedusa Aurelia aurita. We found that flow around swimming ephyrae and their lappets was relatively viscous (1 < Re < 10) and, as a result, ephyral lappets were surrounded by thick, overlapping boundary layers that occluded flow through the gaps between lappets. As medusae grew, their fluid environment became increasingly influenced by inertial forces (10 < Re < 10,000) and, simultaneously, clefts between the lappets were replaced by organic tissue. Hence, although the bell undergoes a structural transition from discontinuous (lappets with gaps) to continuous (solid bell) surfaces during development, all developmental stages maintain functionally continuous paddling surfaces. This developmental pattern enables ephyrae to efficiently allocate tissue to bell diameter increase via lappet growth, while minimizing tissue allocation to inter-lappet spaces that maintain paddle function due to boundary layer overlap.

Unusual development of the mitraria larva in the polychaete Owenia collaris.

Abstract: Despite the wide variety of larval forms among polychaetes, most are clearly derived from the canonical spiralian trochophore. Within the genus Owenia (family Oweniidae), however, the mitraria larva lacks the characteristic ciliary bands of the trochophore, and those it has are monociliated, typically a deuterostome characteristic. Adult Owenia spp. also possess a monociliated epidermis and deuterostome-like nephridia. This study is the first detailed account of early embryology for any member of the Oweniidae. Light, confocal, and scanning electron microscopy were used to investigate organogenesis from fertilization through metamorphosis in Owenia collaris. Equal spiral cleavage yields an embryo with an unusually large blastocoel for a spiralian. The embryo undergoes gastrulation by invagination, and begins swimming 24 h after fertilization. Three important events deviate markedly from stereotypical polychaete embryogenesis. First, at the 8-cell stage the micromeres are larger than the macromeres, as in ...

Larval development with transitory epidermis in Paranemertes peregrina and other hoplonemerteans.

Abstract: We describe development of the hoplonemertean Paranemertes peregrina from fertilization to juvenile, using light, confocal, and electron microscopy. We discovered that the uniformly ciliated lecithotrophic larva of this species has a transitory epidermis, which is gradually replaced by the definitive epidermis during the course of planktonic development. The approximately 90 large multiciliated cleavage-arrested cells of the transitory larval epidermis become separated from each other by intercalating cells of the definitive epidermis, then gradually diminish in size and disappear more or less simultaneously. Rudiments of all major adult structures—the gut, proboscis, cerebral ganglia, lateral nerve cords, and cerebral organs—are already present in 4-day-old larvae. Replacement of the epidermis is the only overt metamorphic transformation of larval tissue; larval structures otherwise prefigure the juvenile body, which is complete in about 10 days at 7–10 °C. Our findings on development of digestive system...

GYRKPPFNGSIFamide (Gly-SIFamide) modulates aggression in the freshwater prawn Macrobrachium rosenbergii.

Abstract: The freshwater prawn Macrobrachium rosenbergii is a tropical crustacean with characteristics similar to those of lobsters and crayfish. Adult males develop through three morphological types—small (SC), yellow (YC), and blue claws (BC)—with each representing a level in the dominance hierarchy of a group, BC males being the most dominant. We are interested in understanding the role played by neuropeptides in the mechanisms underlying aggressive behavior and the establishment of dominance hierarchies in this type of prawn. SIFamides are a family of arthropod peptides recently identified in the central nervous system of insects and crustaceans, where it has been linked to olfaction, sexual behavior, and gut endocrine functions. One of the six SIFamide isoforms, GYRKPPFNGSIFamide (Gly-SIFamide), is highly conserved among decapod crustaceans such as crabs and crayfish. We wanted to determine whether Gly-SIFamide plays a role in modulating aggression and dominant behavior in the prawn. To do this, we performed b...

How did indirect development with planktotrophic larvae evolve

Abstract: The two main types of theories for the evolution of the biphasic life cycles in marine invertebrates are discussed. The "intercalation" theories propose that the larval stages (planktotrophic or lecithotrophic) have evolved as specializations from the ancestral, direct life cycle. The opposing "terminal addition" theories propose that the ancestor was holopelagic and that the adult stage was added to the life cycle with the pelagic stage retained as a planktotrophic larva. It is emphasized that theories based on hypothetical ancestors that were unable to feed must be rejected. This applies to planula theories based on a compact planula. Various arguments against the theories that consider the feeding larvae as ancestral in the major eumetazoan lineages and in particular against the trochaea theory are discussed and found untenable. It is suggested that the "Cambrian explosion" was actually a rapid Ediacaran radiation of the eubilaterians that was made possible by the evolution of a tubular gut with all the resulting possibilities for new body plans.

Pre-settlement behavior in larval bryozoans: the roles of larval age and size.

Abstract: Larval behaviors prior to settlement are impor- tant for both dispersal and the likelihood that larvae will encounter settlement habitat. The role of endogenous fac- tors such as larval age and size are likely to be important in determining pre-settlement behavior but are less well un- derstood than exogenous factors. In a simple experiment we explored the role of larval age and size on pre-settlement behavior in two species of bryozoan. We then used the results of this experiment to develop a theoretical model, which explored potential fitness benefits associated with phenotype-dependent changes in larval behavior (i.e., be- haviors that changed depending on larval age or larval size) in a heterogeneous environment. In the experiment we de- layed the metamorphosis of larvae of Bugula neritina and Watersipora arcuata and assessed the changes in the be- havior of individual larvae (exploring the substratum vs. swimming away from it) as a function of larval age and size. In B. neritina, larval size had no effect on larval swimming behavior, but the youngest and oldest larvae spent more time exploring the substrate than did larvae of intermediate age. In W. arcuata, larval size and age had interactive effects on larval behavior. Our theoretical model predicted that phenotype-dependent behaviors carried a fitness benefit relative to phenotype-independent behaviors, but this de- pended strongly on the availability and quality of habitat elsewhere. We suggest that, taken together, larval age and size are important endogenous factors that act to affect pre-settlement larval behavior and that changes in behavior may act to increase fitness.

The canonical network of autotrophic intermediary metabolism: minimal metabolome of a reductive chemoautotroph.

Abstract: Chemoautorophs that fix carbon by the reductive tricarboxylic acid cycle represent one of the dominant bacterial life forms that make a major contribution to biomass production. From the viewpoint of biogenesis, construction of a canonical chart of intermediary metabolism for this class of organisms may help us to understand early cellular evolution and point us to the last universal common ancestor. Data-mining the KEGG Pathways database enabled us to integrate required biosynthetic pathways and derive a chart that represents the complete anabolic network of a reductive chemoautotroph. Compounds of this metabolic network together constitute a representative minimal metabolome that comprises 287 metabolites. These compounds have been classified into different groups including those compounds that form nodes in the network. It can be seen that a relatively sparse set of organic chemical reactions dominate the anabolic synthesis in the assembly of the minimal autotrophic metabolome. Empirical generalization...

Seawater Temperature Alters Feeding Discrimination by Cold-Temperate but not Subtropical Individuals of an Ectothermic Herbivore

Abstract: Seawater temperature varies across multiple spatial and temporal scales, yet the roles that such variation play in altering biotic interactions are poorly known. We assessed temperature-mediated feeding behavior exhibited by the herbivorous amphipod Ampithoe longimana col- lected from cold-temperate and subtropical estuaries (27°N and 41°N, respectively). Individuals were offered a pairwise feeding choice between lyophilized seaweeds that provide higher fitness (Ulva intestinalis) or lower fitness (Halimeda tuna, H. opuntia, Amphiroa spp., or Stypopodium zonale). Overall, herbivores preferentially consumed the higher quality U. intestinalis more than any lower quality food. However, the strength of this feeding choice was not con- sistent. Northern herbivores consumed proportionally more poorer quality tissue at 25 °C than at 20 °C in two assays (H. opuntia and Amphiroa sp.), consumed less poorer quality tissue at 25 °C than at 20 °C in one assay (S. zonale), and showed no difference in another assay (H. tuna). Moreover, when offered tissue coated with lipophilic extracts of H. opuntia, northern herbivores consumed more extract-coated tissue at 25 °C than at 20 °C. In contrast to northern herbivores, the southern herbivores did not alter their feed- ing choices with temperature. This study represents the first demonstration that short-term (i.e., days-long) variation in ambient temperature alters feeding choices in a marine herbivore.

Cryptic species differentiated in Conus ebraeus, a widespread tropical marine gastropod.

Abstract: Anomalous mitochondrial and nuclear gene sequences in individuals of the widely distributed tropical marine gastropod Conus ebraeus that were not distinguishable by shell shape and color pattern characters suggested the presence of a second, cryptic species. We tested this hypothesis by genetic, morphological, and ecological comparisons of additional individuals from the site in Okinawa where the two forms co-occurred. Radular tooth size and shape, prey type in nature, and microhabitats utilized differed markedly between the two forms. Adults with typical C. ebraeus DNA and radular teeth preyed primarily on errant polychaetes (Eunicidae); those with anomalous DNA and teeth ate mainly sedentary capitellids. Juveniles (shell length <13 mm) had more similar teeth and ate primarily syllids. Radular teeth of the anomalous form agreed with those of Conus judaeus, distinguished from C. ebraeus by Rudolph Bergh in 1895 solely on tooth characters of one specimen from the Philippines. Samples from other widely scat...

Evidence for Costs of Mating and Self-Fertilization in a Simultaneous Hermaphrodite With Hypodermic Insemination, the Opisthobranch Alderia willowi

Abstract: Simultaneous hermaphrodites offer the chance to study antagonistic coevolution between the sexes when individuals function in both roles. Traumatic mating by hypodermic insemination has repeatedly evolved in her- maphroditic taxa, but evidence for the fitness costs of such male-advantage traits is lacking. When reared in isolation, specimens of the sea slug Alderia willowi (Opisthobranchia: Sacoglossa) initially laid clutches of unfertilized eggs but 4 days later began self-fertilizing; this is only the third report of selfing in an opisthobranch. Hypodermic insemination may allow selfing in Alderia if penetration of the body wall bypasses internal mechanisms that promote outcrossing. Selfing specimens and slugs reared in pairs had reduced fecundity compared to isolated slugs laying unfertilized clutches, suggesting that hypodermic insemination imposes a cost of mating. Egg production increased for field-caught slugs separated after mating compared to slugs held in pairs, a further indication that accessibility to mates imposes a fitness cost to the female function. Such antagonism can confer a competitive advantage to slugs mating in the male role but diminish reproduction in the female role among hermaphrodites capable of long-term sperm storage. Alderia willowi is also a rare case of poecilogony, with adults producing either planktotrophic or lecithotrophic larvae. Our rearing studies revealed that most slugs switched be- tween expressed development modes at some point; such reproductive flexibility within individuals is unprecedented, even among poecilogonous species.

Low oxygen consumption and high body content of catch connective tissue contribute to low metabolic rate of sea cucumbers.

Abstract: The energy consumption of echinoderms is low in comparison with that of other invertebrates. We demon- strated this by measuring the oxygen consumption rate per unit of body weight (VO2) of the sea cucumber Actinopyga mauritiana: VO2 was 1/8 that of the "standard" inverte- brates. Low energy consumption in echinoderms has been attributed to their high skeletal content and to catch con- nective tissues (CCTs) that maintain body posture by alter- ing their mechanical properties with little energy expendi- ture. The former is not applicable to holothurians, and the latter has not been proven experimentally. We postulated that the large content of dermal connective tissue, which maintaines posture economically, contributes to the low energy consumption in holothurians. Body-wall dermis oc- cupied 53.5% of wet body weight, whereas body-wall mus- cles, including those of tube feet, occupied 5.1%. VO2 of the dermis in the stiff state (2.45 l g 1 h 1 ) was 1/10 that of the longitudinal body-wall muscle in contraction. The mechanical tests revealed that the stress at an imposed strain of 2% strain was 7 times greater in CCT than in muscles. These results showed that CCT could maintain posture more economically than muscles could. We concluded that the high content of connective tissue with energy-saving pos- ture-maintenance activities contributed to the low energy consumption of holothurians.