Ontogeny of the holothurian larval nervous system: evolution of larval forms

Echinoderm larvae share numerous features of neuroanatomy. However, there are substantial differences in specific aspects of neural structure and ontogeny between the dipleurula-like larvae of asteroids and the pluteus larvae of echinoids. To help identify apomorphic features, we have examined the ontogeny of the dipleurula-like auricularia larva of the sea cucumber, Holothuria atra. Neural precursors arise in the apical ectoderm of gastrulae and appear to originate in bilateral clusters of cells. The cells differentiate without extensive migration, and they align with the developing ciliary bands and begin neurogenesis. Neurites project along the ciliary bands and do not appear to extend beneath either the oral or aboral epidermis. Apical serotonergic cells are associated with the preoral loops of the ciliary bands and do not form a substantial commissure. Paired, tripartite connectives form on either side of the larval mouth that connect the pre-oral, post-oral, and lateral ciliary bands. Holothurian larvae share with hemichordates and bipinnariae a similar organization of the apical organ, suggesting that the more highly structured apical organ of the pluteus is a derived feature. However, the auricularia larva shares with the pluteus larva of echinoids several features of neural ontogeny. Both have a bilateral origin of neural precursors in ectoderm adjacent to presumptive ciliary bands, and the presumptive neurons move only a few cell diameters before undergoing neurogenesis. The development of the holothurian nervous systems suggests that the extensive migration of neural precursors in asteroids is a derived feature. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Drosophila adult and larval pheromones modulate larval food choice

Insects use chemosensory cues to feed and mate. In Drosophila, the effect of pheromones has been extensively investigated in adults, but rarely in larvae. The colonization of natural food sources by Drosophila buzzatii and Drosophila simulans species may depend on species-specific chemical cues left in the food by larvae and adults. We identified such chemicals in both species and measured their influence on larval food preference and puparation behaviour. We also tested compounds that varied between these species: (i) two larval volatile compounds: hydroxy-3-butanone-2 and phenol (predominant in D. simulans and D. buzzatii, respectively), and (ii) adult cuticular hydrocarbons (CHs). Drosophila buzzatii larvae were rapidly attracted to non-CH adult conspecific cues, whereas D. simulans larvae were strongly repulsed by CHs of the two species and also by phenol. Larval cues from both species generally reduced larval attraction and pupariation on food, which was generally—but not always—low, and rarely reflected larval response. As these larval and adult pheromones specifically influence larval food search and the choice of a pupariation site, they may greatly affect the dispersion and survival of Drosophila species in nature.     

Co-option and dissociation in larval origins and evolution: the sea urchin larval gut

SUMMARY The origin of marine invertebrate larvae has been an area of controversy in developmental evolution for over a century. Here, we address the question of whether a pelagic "larval" or benthic "adult" morphology originated first in metazoan lineages by testing the hypothesis that particular gene co-option patterns will be associated with the origin of feeding, indirect developing larval forms. Empirical evidence bearing on this hypothesis is derivable from gene expression studies of the sea urchin larval gut of two closely related but differently developing congenerics, Heliocidaris tuberculata (feeding indirect-developing larva) and H. erythrogramma (nonfeeding direct developer), given two subsidiary hypotheses. (1) If larval gut gene expression in H. tuberculata was co-opted from an ancestral adult expression pattern, then the gut expression pattern will remain in adult H. erythrogramma despite its direct development. (2) Genes expressed in the larval gut of H. tuberculata will not have a coordinated expression pattern in H. erythrogramma larvae due to loss of a functional gut. Five structural genes expressed in the invaginating archenteron of H. tuberculata during gastrulation exhibit substantially different expression patterns in H. erythrogramma with only one remaining endoderm specific. Expression of these genes in the adult of H. erythrogramma and larval gut of H. tuberculata , but not in H. erythrogramma larval endoderm, supports the hypothesis that they first played roles in the formation of adult structures and were subsequently recruited into larval ontogeny during the origin and evolution of feeding planktotrophic deuterostome larvae.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Pathology of larval Eustrongylides in the rabbit

Three fishermen from Maryland who swallowed live bait-minnows developed severe abdominal pain within 24 hr; 2 required abdominal surgery. Larvae of the nematode Eustrongylides sp. were found in the peritoneal cavity of both (Guerin et al., 1982). In the current study, the lesions produced by Eustrongylides larvae were investigated in New Zealand white rabbits. None of these exhibited any signs of clinical illness; however, postmortem examination within 24 hr of inoculation revealed that larvae had migrated through the walls of the esophagus and stomach and viable larvae were recovered from the pleural and peritoneal cavities as well as from gastric contents. Necropsies performed at different intervals of time postinoculation showed that the migrating larvae had produced multi-focal peritonitis and multiple granulomata in the liver.     

Transport of larval fish in the Amazon

In the Amazon near Manaus, larvae of Characiformes, Clupeiformes, Tetraodontiformes, Pleuronectiformes, Gymnotiformes, Belonidae and Sciaenidae were sampled in the river during most of the year, except in June and July, when the water level was at its maximum. Characiformes, Tetraodontiformes and Siluriformes were found in the ichthyoplankton mostly during the rising waters, but Clupeiformes and Sciaenidae drifted in the river almost all year around. Egg abundance was extremely low, suggesting that they do not drift or have a very short residence time. Two types of larval drift seemed to occur: a rising water drift and a lowering water drift. Characiformes, Tetraodontiformes and some Clupeiformes drifted mostly during the rising waters and were more abundant near the banks. Their strategy seemed to be a passive dispersion towards the floodplain with the flood pulse. The high densities near the banks optimized their chances of reaching a floodplain inlet. The groups that drifted during the lowering waters showed an alternative strategy. They were flushed from the floodplain lakes and may have stayed in the main river channel for a few months before returning to the floodplain. Predation in the lakes during the period when water level decline was probably the force behind this drift.     

Modelling the larval dispersal of Acanthaster planci

The desire to understand the observed patterns of Acanthaster population spread throughout the Great Barrier Reef (GBR) has necessitated the development of numerical models which are capable of simulating the hydrodynamics associated with large assemblages of reefs over the time scale of larval dispersal. Models are presented which provide an overview of the physical processes controlling the advection of larvae within the Cairns Section of the GBR Marine Park, on the scale of tens to hundreds of kilometres. The relevant scales and relative importance of the distinct processes are discussed. The models are based on a previously tested numerical hydrodynamic model and are validated by comparison with observations documented in the physical oceanographic literature. Overall, model outputs correspond well to our understanding of the large-scale features and dynamics of water circulation over the continental shelf. Regional variation in circulation predicted by the models may result in significant differences in the movement of larvae.     

Fish larval patterns across the Reykjanes Ridge

Fish larvae and hydrography were surveyed in summer 2005 in the upper 500 m from the Iceland Basin across the Reykjanes Ridge [as part of the Mid-Atlantic Ridge (MAR)], and into the Irminger Sea. Fish larval abundance was highest in the Iceland Basin, followed by the eastern slope of the ridge. Among 22 identified taxa, larvae of mesopelagic boreal and boreal-subarctic species prevailed, with Maurolicus muelleri contributing 80.9% to the total abundance, followed by Benthosema glaciale and Protomyctophum arcticum. Four subtropical species were first recorded for these latitudes, linked with the positive sea surface temperature anomaly encountered. Furthermore, larvae of demersal species were caught, mainly above the ridge and eastern slope of the Greenland Basin, enhancing species richness there. Changes in fish larval abundances, species richness and vertical distributions of larvae coincided with hydrographic fronts and bottom topography. MAR bottom topography guides currents in midwater. The role of these and surface currents for larval dispersal is discussed.     

The effect of larval aggregation behaviour on larval growth of the spruce bark beetle Dendroctonus micans

1. The great spruce bark beetle Dendroctonus micans is a primary pest of spruce in Europe. It is unusual among Eurasian scolytids in that apparently healthy trees are attacked by solitary adults, but larvae feed en masse , in response to a larval aggregation pheromone.
2. The effect of brood size on larval growth was determined in experiments on detached bark, logs and living trees. A positive relationship was found between brood size and larval growth in experiments started with either eggs or larvae up to fifth instar. The relationship appeared to be independent of the effects of both preformed and induced defences in bark.
3. No evidence was found to support the hypothesis that larvae feeding in groups spend a greater proportion of time feeding. Addition of resin to egg chambers in detached bark pieces resulted in high mortality of first-instar larvae. A possible role for larval aggregation in minimizing the effects of host defences is discussed.     

Photoreceptors of the larval tiger salamander retina

Six morphological types of photoreceptor were identified with electron microscopy in radial sections of the retina of the larval tiger salamander, Ambystoma tigrinum. In order of predominance these six types are: red rods, large single cones, double cones composed of principal and accessory members, small single cones, and green rods. The different types of photoreceptor can be distinguished by a number of morphological and cytological characteristics. The identification of the small single cone type now provides evidence for more than one type of single cone in an amphibian retina. The interconnections of the different types of photoreceptor by gap junctions were studied in tangential sections. Rod-rod and rod-cone gap junctions occurred in all possible combinations, but no cone-cone junctions were found even between members of double cones.     

Evolution of larval form in the sea star genus Patiriella: conservation and change in the larval nervous system

The organization of the peptidergic system in the larvae of Patiriella species with divergent ontogenies was compared to determine which aspects of neurogenesis are conserved and which are altered in the evolution of development in these sea stars. P. regularis has ancestral-type feeding bipinnaria and brachiolaria larvae and the organization of the nervous system, in association with feeding structures, paralleled the bilateral larval body plan. P. calcar and P. exigua have non-feeding planktonic and benthic brachiolariae, respectively, and there was no trace of the neuronal architecture involved with feeding. The nervous system in the attachment stage brachiolaria was similar in all three species and neuronal organization reflected larval symmetry. Delayed expression of peptidergic lineages to the brachiolaria stage in the lecithotrophs indicates heterochronic change in the timing of neurogenesis or deletion of the ancestral early neurogenic program. The bipinnarial program is suggested to be a developmental module autonomous from the brachiolar one. With a divergence time of less than 10 Ma, the evolution of development in Patiriella has resulted in extensive reduction in the complexity of the larval nervous system in parallel with simplification in larval form. There is, however, strong conservation in the morphology and neuronal architecture of structures involved with settlement.     

Musca domestica larval lipoprotein

A larval specific high-density lipoprotein (HDL) has been isolated from Musca domestica hemolymph by a combination of density gradient and glycerol gradient ultracentrifugations. The larval lipoprotein has a density of 1.134 g/ml and is formed by at least four apoproteins with molecular weights equal to 26,000, 23,000, 21,000, and 20,000. This lipoprotein contains large amounts of hydrocarbons and phospholipids and minor amounts of diacylglycerols and cholesterol. The larval lipoprotein is completely distinct from lipophorin in regard to apoprotein composition, lipid moiety, physiological pattern, and immunological reactions. Larval lipoprotein is accumulated until the end of the feeding period. During the pupal molt this protein is utilized and is no longer detected after 2 days of pupal stadium. The results obtained imply a possible role of this protein in the puparia and/or pupal cuticle formation. Judging from the properties shown, the Musca domestica larval lipoprotein is a completely new type of insect lipoprotein.     

The effect of larval temperature and photoperiod on the incidence of larval diapause in the blowfly, Calliphora vicina

ABSTRACT. The blowfly, Calliphora vicina Meigen (Diptera: Calliphoridae), displays a maternally induced larval diapause. Progeny of adults exposed to short days enter diapause if the larval temperature is 15C or below; exposing adults to long days, or larvae to temperatures above 15C, results in non-diapause. By keeping progeny from short-day parents at the parental temperature of 23.5C for various lengths of time before transfer to 11C, it could be shown that the diapause-averting effect of high temperature operates in the late wandering stage of larva, possibly at the moment of the decision for the brain-ring gland complex to release PTTH/ecdysone at the diapause/non-diapause stage.
Although photoperiodic induction in the blowfly is maternal, previous work had shown that the larvae were also slightly sensitive to photoperiod. From the present study it is evident that also the eggs are photo-sensitive.     

Advances in the larval rearing of Siberian sturgeon

Since first large-scale attempts to culture sturgeon from the larval stage were carried out in the former U.S.S.R. at the end of the nineteenth century, rearing technology has advanced considerably during the last 20 years and noticeable improvements in incubation and larval rearing techniques have been implemented in normal hatchery procedures. Siberian sturgeon eggs are incubated in MacDonald jars at 13–14° C to prevent fungal infections. Mass hatching takes place 7 days after fertilization and hatching rate can be predicted as a function of the percentage of eggs fertilized. Survival at the end of the endogenous feeding stage is correlated with hatching rate. Egg size has no direct implications for larval growth and survival of Siberian sturgeon. Experimental studies have demonstrated that behavioural observations are useful criteria to assess the quality of larvae and to synchronize the physiological state of fish with the appropriate rearing procedures. Special attention should be given to the transition to exogenous feeding, where cannibalism, difficulties in adaptation to a new diet, overfeeding and resulting bacterial infections dramatically reduces survival to the fingerling stage. Although a commercial artificial diet specifically formulated for larvae of Siberian sturgeon and other acipenserids is still lacking, commercial non-purified rainbow trout diets and starter marine fish diets are currently used and their results are reasonably acceptable in terms of larval growth and survival. Further research must be focused on the determination of egg quality indicators in order to provide the producer with the tools to estimate the viability and performance of theprogeny.