Ovary structure and oogenesis in Cirratulus cirratus (Polychaeta: Cirratulidae)

Oogenesis in polychaetes has, in most species, two main phases: a coelomic phase during which vitellogenesis occurs, and an ovarian phase. Details of the latter are known in very few species. Efficient reproduction requires coordination between these two phases of oocyte differentiation. The ovaries of Cirratulus cirratus (O. F. Müller) occur 2/segment throughout the fertile region of the body; 5 main stages of oocyte differentiation can be recognised on them: stem cells, oogonia, premeiotic oocytes, early oocytes and terminal oocytes. A quantitative study of the composition of the ovary throughout the reproductive cycle shows that the ovary size is approximately constant, but that the percentage of cells in the terminal oocyte stage is reduced to zero prior to spawning. This stage in oocyte differentiation is therefore a critical one, at which oogenesis may be arrested. The ovary alternates between a proliferative phase when the terminal oocytes are present, and a non-proliferative phase when they are absent. The overall rate of germ cell production for each ovary is low, less than 1 cell/ovary/day, and the, ovary tissue turnover time is greater than 2 years. A qualitative model describing the production of oocytes is presented, and the implications of the low rate of germ cell production are discussed. A low rate of germ cell proliferation, as in Cirratulus cirratus, implies that the rate of oocyte output into the coelom will be controlled by factors influencing the later stages of ovarian oocyte differentiation.     

Ultrastructure of sperm and spermathecae in Micromaldane spp. (Polychaeta: Capitellida: Maldanidae)

Assessing the possibility that external fertilization has re-evolved requires the study of monophyletic groups that exhibit various reproductive methods. Maldanid polychaetes show a range of reproductive mechanisms, though previous studies of reproduction have hitherto been restricted to larger species with external fertilization. Micromaldane pamelae Rouse and M. nutricula Rouse are small, gonochoristic maldanids that brood directly developing larvae. Both species have sperm with elongate nuclei and an acrosome extending down each side of the anterior end of the nucleus. A true midpiece is absent; two mitochondria extend along the posterior region of the nucleus. Spermatids develop synchronously in large clusters connected by a cytophore. In M. pamelae sperm are released into the water as spermatozeugmata. These are comprised of clusters of sperm with their tails oriented to the centre and the sperm heads facing outwards. Females of M. pamelae and M. nutricula bear pairs of spermathecae ventrally (M. pamelae three pairs, between setigers 10 and 11, 11 and 12 and 12 and 13 and M. nutricula two pairs, between setigers 10 and 11 and setigers 11 and 12). The blind sacs are epidermal invaginations bound closely together. The entrance to each spermatheca may only be 1 to 2 m across with each spermatheca holding several hundred sperm. This represents the first detailed study of spermathecae in the Capitellida. The occurrence and structure of spermathecae and spermatozeugmata in other groups are discussed and compared with Micromaldane spp. Comparisons are made with non-polychaetes with the purpose of discussing functional aspects of reproductive mechanisms in marine metazoans in general. Elongate sperm nuclei are associated with sperm storage and/or large egg size. The lack of an elongate sperm midpiece may be an indicator of having to swim in water but does not contraindicate sperm storage. Spermatozeugmata may serve as an indication of sperm storage and brooding of larvae. Speculations on the phylogenetic significance of these reproductive features are limited by the fact that supposedly modified (i.e., derived) states may reflect functional/structura, constraints of small body size.     

Origin and development of the amoebocytes of Nicolea zostericola (Polychaeta: Terebellidae) with a discussion of their possible role in oogenesis

Three types of amoebocytes are present at different stages in the life history of Nicolea zostericola (Grube, 1860): a granular amoebocyte or spindle cell and 2 agranular amoebocytes, designated Type I and Type II. The agranular amoebocytes originate from specific sites on the lateral parietal peritoneum of the coelomic cavity in all the thoracic and a few abdominal segments. The origin of the granular amoebocyte is unknown. In juvenile worms the Type I amoebocyte enters the coelomic fluid and accumulates lipid and glycogen, presumably derived from reserves stored in the parietal peritoneum. Near the beginning of the vitellogenic phase of oogenesis, the Type I amoebocyte disappears and a Type II form is produced by the peritoneum. Type II cells differ cytologically from the Type I form by containing extensive rough ER (endoplasmic reticulum), Golgi complexes and free ribosomes; they appear to synthesize protein for export. At the end of vitellogenesis and near spawning, amoebocytes undergo alterations which include the migration of the nucleus to a central position, the formation of stacks and whirls of rough ER cisternae, and the appearance of vacuoles throughout the cytoplasm. It is suggested that the agranular amoebocytes obtain nutrients from food reserves stored in the parietal peritoneum and transfer them to the oocytes for use in yolk synthesis. The function of the granular amoebocytes is unknown.Contribution No.37 from the Marine Science Institute, Northeastern University, Nahant, Massachusetts, USA.Contribution No. 50, Harbor Branch Foundation, Inc.     

Ultrastructure and behavior of the larva of Phragmatopoma californica (Polychaeta: Sabellariidae): identification of sensory organs potentially involved in substrate selection

The tentacles of the larvae of Phragmatopoma californica (Fewkes) a tubicolous, reef-building polychaete, were examined by video-equipped light microscopy and transmission and scanning electron microscopy. The surface of the tentacles has a unique ciliation pattern, consisting of dorsal tufts of short immotile cilia, ventrolateral tufts of short and long immotile cilia, and ventral motile cilia. Cells bearing immotile cilia are primary sensory cells with long basal processes that form synapses with basiepithelial nerve fibers. The sensory cell cytoplasm is similar to that of nervous tissue, and contains microtubules, neurofilaments, and synaptic vesicles. Sensory cell synapses with basiepithelial nerves appear to be both axodendritic and axoaxonic. The structure of the immotile cilia is compared to that of motile cilia. Unlike motile cilia, immotile cilia are short, rigid, end in a blunt tip and possess and axoneme with typically arranged mictotubules that terminate in an electron-dense end plate. The basal feet of immotile cilia do not anastomose with adjacent basal bodies, and the ciliary membrane is loosely applied to the axoneme and is covered by a surface coat of filamentous material. The use of the larval tentacles during substrate exploration, and the location and ultrastructure of sensory cilia, indicate that they may be involved in the perception of substrateassociated chemical signals and/or mechanical cues of significance in substrate selection.     

Gametogenesis in Harmothoe imbricata (Polychaeta: Polynoidae)

The processes of gametogenesis in Harmothoe imbricata are described and illustrated. For most of their development the gametes are attached to the walls of segmental blood vessels. In males, only meiosis and sperm differentiation occur while the cells are free in the coelom. In females, oocytes are only released into the coelom briefly, after vitellogenesis has been completed. H. imbricata thus differs from most polychaete species so far described — where in males the spermatogonia divide mitotically to form rosettes after they have been released into the coelom, and in females the whole of vitellogenesis occurs after release of oocytes into the coelom. The oocytes of H. imbricata enter the first metaphase of meiosis before they are shed at spawning. The elongated head of the sperm in H. imbricata is atypical of polychaetes in general.     

Ultrastructure and histochemistry of the integument of air-breathing polychaetes from mangrove swamps of sumatra

The integument of S.E. Asian air-breathing polychaetes exhibits a variable ultrastructure with regard to cuticle and epithelial cells. The cuticle varies in thickness, contents of regularly arranged collagen fibrils, and structure of the epicuticle. The basal portion of the epithelium is either of smooth outline, or consists of slender processes of the epidermal cells separated by wide intercellular spaces which are invaded by a loose system of glia cells and nerve fibres. Extracellular spaces within the epidermis of the gills of Dendronereides heteropoda are transformed into blood lacunae. The enzyme content of the epithelial cells is relatively low.Supported by the Deutsche Forschungsgemeinschaft (Sto 75/2, We 380/4).     

Anchorage of embryos in Spirorbinae (Polychaeta)

Methods of incubation are taxonomically important in Spirorbinae. About half of the known species incubate in the operculum, but the remainder are usually described simply as incubating within the tube. Studies of tube incubation show that the embryos are anchored in various ways. They form an egg string, which is attached to the tube by a posterior filament in Spirorbis and Spirorbella; they adhere more generally to the tube wall in Circies and Paradexiospira; they are attached to specialised thoracic stalks in Protolaeospira, Helicosiphon and Romanchella; they adhere to the body surface in Metalaeospira and probably in Eulaeospira; they are not attached to either body or tube in Paralaeospira. Tube incubation is found predominantly in temperate or cold waters. In most species from the southern hemisphere, embryos are anchored to the bodies of the parents, but in most species from the northern hemisphere, embryos are fixed to the walls of the parent tubes.     

Tube-worm-sediment relationships of Diopatra cuprea (Polychaeta: Onuphidae)

Diopatra cuprea (Bosc) builds a long, nearly vertical parchment tube in sediments, which is reinforced with bits of shell, sediment or debris. The tube can be divided on the basis of structure into reinforced sections built at or above the sediment surface, and unreinforced sections built below the sediment surface. If sediment accumulates faster than the worm can build reinforced tube, an unreinforced section results. When this occurs, D. cuprea builds reinforced tube at the new sediment surface, and then pulls it down into the substratum until it joins the next lower reinforced section, or until the worm can no longer overcome sediment resistance. In the latter case, the unreinforced section occurs between two reinforced sections. The tube cap is trimmed as sediment is eroded from around it. Selection for reinforcing materials occurs only in the transitional zone from unreinforced to reinforced tube. The exposed portion of the tube (the tube cap) extends from 1 to 6 cm above the sediment surface as an inverted J. The tube mouth is oriented perpendicularly to the direction of current flow. Tube maintenance continues at temperatures below feeding-response blockage (5° to 10°C), but ceases below about 1.8°C.     

Fecundity and reproductive effort in Ophryotrocha labronica (Polychaeta: Dorvilleidae)

Reproductive effort in terms of fecundity and energy allocation was studied in Ophryotrocha labronica La Greca and Bacci, 1962 a small, semicontinuous iteroparous species. In O. labronica fecundity is more or less constant throughout life, and the total fecundity of the 64 couples examined in this experiment only declined by 12% from the first to the last spawning. No linear relationship was found between fecundity and body size.␣The energy content of germinal and somatic tissues was determined by differential scanning calorimeter. The␣reproductive effort and a reproductive index based on the fifth spawning were evaluated to compare the␣patterns of energy allocation of a semicontinuous iteroparous species with semelparous and annual iteroparous species. The reproductive index ranged from 0.20 to 0.77, with an average of 0.480. The average reproductive effort was 0.840, ranging from 0.60 to 0.96. These figures highlight the enormous amount of energy that O. labronica allocates to reproductive tissues. Received: 23 July 1997 / Accepted: 20 October 1998     

Hemolysins from the mucus of Spirographis spallanzani (Polychaeta: Sabellidae)

Specimens of Spirographis spallanzani (Polychaeta: Sabellidae) were collected in the Gulf of Taranto, Italy, in 1991, and their mucous secretion was examined. The mucus possesses thermolabile biologically active proteins which produce hemolysis in vertebrate erythrocytes. The molecules are ineffective against both unfertilized and fertilized echinoid eggs and spermatozoa. The lytic reaction against rabbit erythrocytes varies as a function of cations, pH and temperature, and displays a fast time-course. No immunological cross-reaction could be demonstrated between hemolysins of S. spallanzani and Eisenia foetida andrei. SDS-PAGE analysis of mucus proteins reacting with sheep erythrocytes revealed that at least three major (16, 22 and 40 kDa) and two minor (29 and 34 kDa) components interact with membrane targets.     

Secondary production ofPrionospio caspersi (Annelida: Polychaeta: Spionidae)

The annual somatic production ofPrionospio caspersi Laubier was estimated, between July 1986 and 1987, at a sandy shallow-bottom station in the area facing the Po River Delta, Italy, where this species is one of the most abundant. Primary recruitment occurs in summer, and maturation of gametes the following spring. Body sizes were estimated from the surface area of the anterior end of worms, projected onto a monitor and recorded using a digitizer tablet. Measurements, to the tenth setiger, were correlated with total length, total body surface area and AFDW. The annual production was 8.06 g m^–2 yr^–1 (AFDW) and the production:biomass (P:B) ratio 4.09. The results are discussed in relation to some physical and chemical features of the environment, stressing differences with the life cycle of other coexisting populations.     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

Evolution of reproductive features and larval development in the genus Amphiglena (Polychaeta: Sabellidae)

Larval development in Amphiglena mediterranea is direct in the parental tube, and larvae crawl away as young juveniles with a radiolar crown capable of feeding. The non-feeding larvae of A. nathae are initially brooded in the parental tube, and then have a swimming phase before settling and developing a radiolar crown. Developmental data and reproductive features such as sperm structure are combined with other morphological data for an assessment of the cladistic relationships of the species in the genus Amphiglena. Two species of Laonome are included as part of the ingroup. Six genera, representing a sister group to the Laonome/Amphiglena clade, are used as an outgroup. Two most-parsimonious cladograms were found, and the evolution of reproductive features are discussed. The monophyly of Amphiglena is indicated by features previously identified, as well as two features based on sperm morphology. At present a plesiomorphic larval form cannot be identified for the genus Amphiglena. Received: 16 June 1997 / Accepted: 29 January 1998     

Population dynamics and secondary production in Melinna palmata (Polychaeta: Ampharetidae)

A Southampton Water (England) population of Melinna palmata Malmgren was studied from July 1978 to March 1980. M. palmata is a gregarious sedentary polychaete living in muddy substratum. The distribution of the polychaete in Southampton Water was related to the silt content of the sediment. The species occurs in sediments of <60% silt, it is dioecious, and the gametes are shed during a prolonged breeding season throughout the year, with peaks in March and July. Mature oocytes measure between 180 and 240 m in diameter at spawning, and fertilization is external. Unshed oocytes are resorbed. Annual secondary ary production was 0.42 gC m^–2yr^–1 and the P:B ratio was 2.19.     

Suspension-feeding and assimilation efficiency in Lanice conchilega (Polychaeta)

In laboratory experiments, measurements of ingested ration, assimilated ration and assimilation efficiency in the polychaete Lanice conchilega (Pallas) were carried out at 12°C and at a food concentration of 40x10⁶ cells of Dunaliella marina/l. Over periods of 20 to 46 days, the ingested ration was determined photometrically by the continous automatic recording apparatus described by Winter (1973). The assimilated ration was calculated from the calorific content of the food ingested minus the calorific content of the faeces produced. Within the range of body size investigated (1.3 to 35.0 mg dry-tissue weight), the food intake (=filtration rate=F) was found to be a linear function of body size (W) according to the equation F=0.88 W ^0.32. The daily food intake-expressed as percentage of dry-tissue weight-decreased from 35.7 to 3.9% with increasing body size. The assimilation efficiency was found to vary between 70.6 and 77.2% and showed no significant correlation with body size. The absolute amounts of food retained from suspension and the assimilation efficiencies calculated for L. conchilega are within the range typical for obligatory suspension-feeding organisms. This implies that L. conchilega is capable of completely replacing deposit-feeding by suspension-feeding.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung-Abwässer in Küstennähe.     

Functional morphology and feeding behavior of Paraprionospio pinnata (Polychaeta: Spionidae)

The functional morphology and feeding behavior of Paraprionospio pinnata (Ehlers) were studied. The tentacular feeding palps of P. pinnata possess four groups of cilia: laterals, latero-frontal cirri, frontals and basal transverse rows. The lateral cilia beat in metachronal waves creating in current that flows toward the frontal surface of the palp. The latero-frontal cirri deflect suspended particles onto the frontal surface and potentially resuspend deposited particles. The frontal cilia line the groove of the palp and transport particles to the ciliated pharynx. The basal transverse cilia also beat in metachronal waves and together with the frontal cilia of the branchiae produce a U-shaped current that removes particles rejected by the pharynx from the burrow. The significance of particle selection at the site of the pharynx in spionid polychaetes is hypothesized to be constrained by the interaction of specific morphologies and behaviors that reduce the effects of epifaunal predation and browsing. The relationship between sediment permeability and type of respiratory or ventilation current is hypothesized to be a potentially important factor in understanding the distribution of spionid polychaetes.     

Life history and population dynamics ofKinbergonuphis simoni (Polychaeta: Onuphidae)

The relationship between life-history characteristics and population dynamics were investigated in the onuphid polychaeteKinbergonuphis simoni (Santos, Day and Rice) between 1982 and 1987. The studied population is located in Upper Tampa Bay, Florida, USA. This worm attains sexual maturity at 5 to 10 mo of age, depending on temperature. Several consecutive broods are produced during a female's life time, with 7 to 26 young per brood. Adults die in June–July after the breeding season. Generations of breeding individuals do not overlap and the replacement of generations occurs in July–August. Only one extended breeding season is experienced per life time. Life span does not exceed 2 yr. Field population density in 1982 was high in fall and spring (2000 to 9000 individuals/m²), low in summer (1600 individuals/m²), and slightly depressed in winter (2500 to 4000 individuals/m²). Changes in population density may be explained by seasonal, temperature-controlled changes in instantaneous birth and death rates. Birth rates are high in fall and spring, while death rates are high in early summer. Death of juveniles contributes to the winter density decrease.     

Zur sexuellen Differenzierung von Ophryotrocha puerilis (Polychaeta: Eunicidae)

Intact young males of Ophryotrocha puerilis Clap. Mecz. which are about to reach the female phase are capable of producing oocytes even under conditions of fasting. Intact isolated females may change sex under fasting conditions; their oocytes are not shed but resorbed. Decerebrated males which are about to reach the female phase at the moment of decerebration are unable to develop oocytes. Decerebrated females begin earlier with the production of spermatozoa than females under fasting conditions. The oocytes of decapitated females are either resorbed (young females with oocytes ranging in diameter up to 60 μm), or shed in egg masses capable of development after fertilization (older females with oocytes of 90 μm diameter or more). The effect of decerebration on females is compensated completely by implantation of isolated prostomia from female donors. Decerebrated males with implanted isolated prostomia from female donors immediately start oocyte production; these oocytes are not shed. In the majority of cases, decerebrated females change to the male phase after implantation of prostomia isolated from male donors. The mutual influence of a pair depends on direct contact of the partners. In isolated females, anterior fragments, consisting of prostomia and some segments, may induce change of sex and shedding of oocytes. On the other hand, intact females may induce sex reversal in anterior fragments. Even isolated prostomia from female donors may induce change of sex in intact females. A possible influence of intact females upon isolated prostomia was not found. Intact females and decerebrated females with implanted prostomia from female donors exhibit no mutual influence when placed together.     

Rates of seasonal sediment reworking in Axiothella rubrocincta (Polychaeta: Maldanidae)

Sediment reworking rates of Axiothella rubrocincta (Johnson, 1901) (Polychaeta: Maldanidae) were measured in situ in Tomales Bay, California (USA), from August, 1969 through July, 1970. On the average, each adult worm (approximate fresh weight=1 g) reworks about 5 g dry sediment d^-1 at a mean temperature of 13.4°C and a mean salinity of 31.8.Reworking rates are positively correlated with temperature and salinity, and negatively with sediment organic carbon, sedimentation rates and grain size. An inverse correlation exists between sediment reworking rates (g dry sediment g^-1 wet wt of worm d^-1) and g wet weight of worm. Sediment parameters deseribing unworked sediments are not significantly different from those for fecal sediments. Although these data suggest this species to be a non-selective deposit-feeder, it is more likely that it is faculatively selective.     

Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae)

Functional morphology and feeding behavior of Scolelepis squamata (Müller) were studied. Gut contents consisted of unconsolidated sedimentary particles, fecal pellets of other species, and a wide variety of embryos, larvae, and juveniles. Unlike other spionid polychaetes the palps of S. squamata lack a median, ciliated groove. Particles captured by the palps were brought to the pharynx by a complete contraction of the palp. In the presence of a current, S. squamata helically coiled their palps, and fed almost exclusively on suspended and resuspended particles. In contrast to most spionid polychaetes the palps of S. squamata are not deciduous. Various aspects of the morphology and feeding behavior of S. squamata are discussed in terms of their functional and ecological significance.