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Phylogenetic analyses of mode of larval development.

Phylogenies based on morphological or molecular characters have been used to provide an evolutionary context for analysis of larval evolution. Studies of gastropods, bivalves, tunicates, sea stars, sea urchins, and polychaetes have revealed massive parallel evolution of similar larval forms. Some of these studies were designed to test, and have rejected, the species selection hypothesis for evolutionary trends in the frequency of derived larvae or life history traits. However, the lack of well supported models of larval character evolution leave some doubt about the quality of inferences of larval evolution from phylogenies of living taxa. Better models based on maximum likelihood methods and known prior probabilities of larval character state changes will improve our understanding of the history of larval evolution.

Animals↗

A statistical analysis of Neanthes arenaceodentata, sister chromatid exchange data.

This paper presents a statistical evaluation of the sister chromatid exchange (SCE) assay using Neanthes arenaceodentata. Specific issues discussed are distribution of baseline SCE frequencies, transformations of SCE data, statistical sensitivity of the assay, repeatability of test results, and sources of variability of SCE data. Probabilities of false positives and false negatives are considered and recommendations are made for appropriate transformation of data, experimental design, and test conditions.

Analysis of Variance↗

Characterization of connectin-like proteins of obliquely striated muscle of a polychaete (Annelida).

In obliquely striated muscle of polychaete, Neanthes sp., three kinds of connectin (titin)-like high molecular weight proteins, approximately 4000 kDa, approximately 1200 kDa and approximately 700 kDa, were detected by SDS gel electrophoresis and immunoblots using antibodies to vertebrate skeletal muscle connectin and antiserum to the protein in question. The 700 kDa protein was isolated and characterized as a beta sheet-rich filament 170 nm long and 4 nm wide. Using polyclonal antibodies to the 700 kDa protein, the binding of the immunogold to the thick filament was only demonstrated in high ionic strength relaxing solution which solubilized some myosin. This observation suggested that the 700 kDa protein was localized below the layers of myosin in the thick filament and this localization is different from that of twitchin of C elegans bodywall muscle that is on the surface of thick filament. The 4000 kDa protein was identified as a very thin filament linking the thick filament to the dense body. The very thin filaments were visualized in gelsolin-treated actin filament-free fibres. The 1200 kDa protein was located in the periphery of the dense body. A model of the elastic filament in polychaete bodywall muscle is presented.

Actin Cytoskeleton↗

Molecular phylogeny of some polychaete annelids: an initial approach to the Atlantic-Mediterranean speciation problem.

The polychaete Eupolymnia nebulosa (family Terebellidae) displays two alternative modes of reproduction. In the Mediterranean, larvae are brooded in a mucous mass while in the Atlantic and English Channel, larvae follow a plank-tonic development. This paper attempts to discern whether this difference is expressed at the population, infraspecies, or species level. Specimens of E. nebulosa and representatives of a number of control species were sampled from Atlantic/English Channel and Mediterranean locations. Genetic sequencing of the Large-subunit ribosomal RNA 5' end of six representative species allowed one to infer the relative position of E. nebulosa within the Terebellidae and the position of the latter within the animal kingdom. The relative genetic distances calculated between the different species were also used to approach the speciation problem raised by the differences between the Mediterranean and Atlantic/English Channel population of E. nebulosa. The genetic distance between Mediterranean and Atlantic populations of both E. nebulosa and Lanice conchilega are of the same order, suggesting that differences between the populations of E. nebulosa are infraspecific.

Animals↗

Evolutionary optimization and neural network models of behavior.

One of the main challenges to the adaptionist program in general and the use of optimization models in behavioral and evolutionary ecology, in particular, is that organisms are so constrained by ontogeny and phylogeny that they may not be able to attain optimal solutions, however those are defined. This paper responds to the challenge through the comparison of optimality and neural network models for the behavior of an individual polychaete worm. The evolutionary optimization model is used to compute behaviors (movement in and out of a tube) that maximize a measure of Darwinian fitness based on individual survival and reproduction. The neural network involves motor, sensory, energetic reserve and clock neuronal groups. Ontogeny of the neural network is the change of connections of a single individual in response to its experiences in the environment. Evolution of the neural network is the natural selection of initial values of connections between groups and learning rules for changing connections. Taken together, these can be viewed as "design parameters". The best neural networks have fitnesses between 85% and 99% of the fitness of the evolutionary optimization model. More complicated models for polychaete worms are discussed. Formulation of a neural network model for host acceptance decisions by tephritid fruit flies leads to predictions about the neurobiology of the flies. The general conclusion is that neural networks appear to be sufficiently rich and plastic that even weak evolution of design parameters may be sufficient for organisms to achieve behaviors that give fitnesses close to the evolutionary optimal fitness, particularly if the behaviors are relatively simple.

Animals↗

Possible involvement of putrescine in nucleolar formation in early embryos.

Continuous treatment of developing eggs of the polychete Ophryotrocha labronica with alpha-methylornithine, which inhibits synthesis of putrescine, led to arrest of development at gastrulation. The present ultrastructural analysis suggests that the arrest of development is due to failure to form nuclei, and thus reveals a possible role for putrescine in nucleolar formation. Further support for this contention was provided by means of electron-microscopical autoradiography. It was found that newly synthesized putrescine, derived from administered 3H-ornithine, labeled the nucleoli intensely at the time of their normal appearance during gastrulation, the time at which the rate of endogenous putrescine synthesis is maximal. These observations have led to the conclusion that putrescine synthesis may be directly involved in formation of nucleoli.

Animals↗

Modified axonemes and ciliary membranes in three polychaete species.

In living Ophryotrocha puerilis, Polyophthalmus pictus and Dinophilus gyrociliatus no modified cilia are present. Treatment with hyper- and hypotonic magnesium chloride solutions leads to the formation of either cilia with dilated tips or discocilia (paddle cilia). Discocilia show axoneme loops within distal swellings of the ciliary membranes. Both types of modified cilia regain their normal appearance if they are allowed to recover in seawater. The total number of discocilia and the diameter of the loops are inversely related to the osmolarity of the magnesium chloride solution used. Even isotonic solutions of magnesium chloride, which are usually used to anaesthetize marine worms, readily induce modified cilia. This indicates that the effect is not merely due to osmotic conditions. Glutaraldehyde and osmium tetroxide may act in the same way to induce modified cilia, a fact which may account for the numerous TEM and SEM documentations of modified cilia in various marine invertebrates. Which cilia in a particular species are modified varies from one specimen to another.

Animals↗

The structure of symplasmic early oocytes and their enveloping sheath cells in the polychaete, Platynereis dumerilii.

1. Early oocytes of Platynereis dumerilii are found in clusters floating in the coelom. The oocytes of a cluster form a syncytium which is enveloped by several sheath cells. 2. At stage 2, only a single sheath cell per cluster remains, and it penetrates the group of rounded oocytes, enveloping each one of them. At stage 4, this cell contains a reticular basket made up of bundles of filaments and is inferred to be degenerating, from the presence of vacuoles, clumps of pigment-like material, and atypical mitochondria. 3. Synaptonemal complexes are typical of the nuclei of stage 2 oocytes. Oocytes of stage 4 (early vitellogenesis) contain stacks of endoplasmic reticulum in a distinctive arrangement, with interspersed electron-dense masses. Similar masses accumulate in the cytoplasm close to the nucleus and adjacent to the nuclear pores. 4. From the present observations, a physical supporting rather than a nutritive function is attributed to the sheath cell, which ensures cohesion among the oocytes connected with each other throughout the cluster phase by cytoplasmic bridges. This finding is discussed with respect to conclusions drawn from oocyte transplantation experiments.

Animals↗

The fine structure of polychaete septate junctions.

Epidermal septate junctions of Nereis sp. and Cirriformia sp. fixed with OsO4 or glutaraldehyde/OsO4 display variable structure in electron micrographs. In transverse section the septa are often indistinct and obscured by opaque material that fills the junctional cleft. Septa (spaced at 180--280 A) are more clearly defined in slightly oblique transverse section; they exhibit an electron lucent center and appear to be linked by arms. En face views of the junction show a honeycomb pattern. Cytoplasmic faces of junctional membranes are backed with plaques opposite the septa. Lanthanum used as a tracer delineates junctional structure in negative contrast. In transverse section a chain-like lattice is present in the junctional cleft. En face vies show parallel rows of pleated elements often linked by arms into honeycomb arrays. Oblique sections demonstrate that these pleated elements are continuous with the chain-like lattice seen in transverse sections. Lanthanum does not pass entirely through the junction. Lanthanum reveals that the septa have a very intricate substructure, but it is difficult to visualize the architecture that could generate the various images presented by these junctions when seen in different orientations. However, it is clear that these junctions possess some features that are diagnostic of several supposedly different types of septate junctions in invertebrates.

Animals↗

Effects of light and dark on photoreceptors in the polychaete annelid Nereis limnicola.

The effects of light and dark on photoreceptors of the brackish-water polychaete annelid Nereis limnicola were studied by electron microscopy. Animals dark-adapted for one or two days exhibited well-formed straight microvilli (rhabdomeres) on the sensory cell processes. Continuous illumination of worms for one or two days caused extensive breakdown of the microvilli into vesicles and debris. Thirty minutes to three h of exposure of dark-adapted animals to light produced increasing severity of degradation of photoreceptoral microvilli. Light-adapted worms placed in darkness for one-half to three h showed progressive restoration of the microvilli to the dark-adapted condition. The products of degradation were internalized by both sensory and pigmented supportive cells by phagocytosis and pinocytosis.

Animals↗

Cytochemical localization of acid phosphatase in light- and dark-adapted eyes of a polychaete worm, Nereis limnicola.

The amount and distribution of the lysosomal enzyme acid phosphatase in light- and dark-adapted eyes of the brackish-water annelid Nereis limnicola were studied by standard cytochemical techniques. Precipitate from the acid phosphatase reaction was observed in Golgi-endoplasmic reticulum-lysosomal complexes, primary lysosomes, and secondary lysosomes, formed by fusion of primary lysosomes with phagocytic and pinocytic vesicles containing products of presumed rhabdomeric degradation. The acid phosphatase reaction occurred in these organelles in both sensory and supportive cells of both light- and dark-adapted ocelli. Secondary lysosomes were more abundant in sensory cells of illuminated ocelli than in those maintained in the dark. Sparse reaction product was found in Golgi cisternae, none in rough endoplasmic reticulum. We suggest that the increase of lysosomal activity in light-adapted eyes is correlated with the breakdown of photosensory microvilli upon exposure to light. A diagram of our interpretation of recycling of photoreceptoral membrane in N. limnicola is presented.

Acclimatization↗

Morphogenesis of larval cuticle in the polychaete Phragmatopoma lapidosa. A correlated scanning and transmission electron microscopic study from egg envelope formation to larval metamorphosis.

The development of the egg envelope and its incorporation into the larval cuticle of the polychaete Phragmatopoma lapidosa, was studied by correlative scanning and transmission electron microscopy. The mature egg possesses an envelope composed of five zones including an outer granular zone formed by the tips of the egg microvilli. The formation of the granules is described and their functions are discussed. The entire egg envelope is retained as the larval cuticle up to the 16 h trochophore stage. From this stage to about the 60 h larval stage, the envelope is gradually lost and replaced by a cuticle consisting of branching microvilli. The cuticle of the 20 day larva is composed of highly branching microvilli penetrating a homogeneous electron opaque cuticle. The possible functions of the cuticle among the Annelida are discussed.

Animals↗

A scanning electron microscopic study of larval development in the marine polychaete, Galeolaria caespitosa Lamarck (Serpulidae).

Gametes and developing larvae of the polychaete Galeolaria caespitosa were examined by scanning electron microscopy. The sperm display a primitive morphology. When treated with 0.33 M CaCl2, they release a branched acrosomal process. At spawning, the polygonal oocytes have a granular surface made up of spherules and the tips of microvilli. The oocyte coat develops a ridged appearance as the oocyte rounds up. At fertilization, the microvilli are withdrawn from the coat surface. Microvilli again appear on the coat surface during the trochophore stage, but the egg coat appears to be retained as the larval cuticle until the demersal stage. The surface of the larva now shows many microvilli. Details of the organization of several ciliary structures are clarified. Moreover, the present study shows rapid, sequential development of paired setal sacs, with the most anterior pair appearing first.

Animals↗

Sodium channel opening as a precursor to inactivation. A route to the inactivated state.

The time constant of the process producing the delay in Na inactivation development as determined by the two pulse method (tau delay) was extracted and compared to that of the slowest Na activation process (tau 3) for the INa during the conditioning pulse of that same determination. tau delay and two pulse inactivation (tau C) values were computer generated using a non-linear least squares algorithm. tau 3 and single pulse inactivation (tau h) values were independently generated for each determination also with the aid of the computer using the same non-linear least squares algorithm. In one determination at 2 mV, tau C was 4.68 and tau delay 0.494 ms while tau h was 4.70 and tau 3 0.491 ms for a tau c/tau h of 0.996 and a tau delay/tau 3 of 1.006. Mean tau delay/tau 3 from five determinations in four axons, both Cs and K perfused, and spanning a potential range of -27 to 2 mV was 1.068. The precursor process to inactivation is channel opening. Some fraction of channels presumably inactivate via another route where prior channel opening is not required.

Animals↗