Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

Reproductive cycle of the subarctic brooding asteroid Leptasterias polaris

The reproductive cycle of the large brooding seastar Leptasterias polaris Müller and Troschel was examined over an 18-month period in the St. Lawrence Estuary, Québec, Canada. There is a distinct annual cycle with spawning over several months in the autumn. The female has an unusual cycle in that the ovary only slightly decreases in size during spawning, and the size structure of the oocyte population is remarkably stable throughout the year. The major annual change observed in the oocyte population is the development of a small number of 600-to 800- oocytes prior to spawning and their loss during spawning. This stability, combined with the lack of evidence of phagocytosis, suggests that oocyte development takes place over many years. By contrast, the developmental cycle of the testis is similar to that of most echinoderms. The large reservoir of oocytes probably guarantees a steady annual recruitment, and brooding during the winter probably minimizes metabolic costs for the female and ensures the release of the juveniles when conditions are favourable in the spring and summer.     

Sexual reproduction and settlement of the coral reef spongeChalinula sp. from the Red Sea

Characteristics of the sexual reproduction and larval settlement of the haplosclerid spongeChalinula sp., which inhabits the shallow waters (1 to 6 m) of Eilat, Red Sea, were investigated from September 1985 through to November 1987. This species was found to be a simultaneous hermaphroditic brooder, hence gonochorism is not the rule in the order Haplosclerida. Brooding always takes place in special brooding chambers. While the oocytes in the brooding chambers are among the largest known in sponges (355±37µm), the spermatic cysts distributed in the choanosome are among the smallest known for this phylum (average 26±7µm).Chalinula sp. breeds throughout the year and in experiments most larvae (74%) settled within 1 to 8 h post-release, generally within 4.5 h. Metamorphosis from larval shape to a sessile sponge lasts 1 to 6 h. Thus, larvae had a short swimming period, settled fast, and metamorphosed rapidly (within 1 to 6 h). The large size of the larvae may contribute to their ability to rapidly reorganize their body shape into that of a sessile sponge. In addition, the existence of already differentiated choanocyte chambers in the larvae, facilitates fast construction of the water filtration system in the newly settled sponges. The reproductive and larval characteristics ofChalinula sp. enable the larvae to settle on any vacant space in the reef, which may explain its abundance in the Red Sea.     

Sex in the tropics: reproduction of<Emphasis Type="Italic"> Chthamalus malayensis</Emphasis> Pilsbury (Class Cirripedia) at the equator

The reproduction of a key intertidal space occupier, the barnacle, Chthamalus malayensis, was examined over a 13-month period. The reproductive cycle of C. malayensis was studied at three shores (Singapore, east and west coast of Malaysia) near the centre of its geographical range. Testes, vesiculae seminales, ovaries and embryos were scored for their stages of development. Although, in general, C. malayensis contained gonads throughout the year, the timing and level of peak development varied between the three shores. Embryos were present throughout the year in west Malaysia but were absent for 4–5 months in east Malaysia (June–August and October) and Singapore (February, April and August). Variations in the reproductive cycles and brooding were not clearly linked to the timing of monsoons. The lack of seasonal peaks in breeding may be attributed to the relatively constant climatic conditions experienced near the equator.Communicated by M. S. Johnson, Crawley     

Reproduction in the sea pen <Emphasis Type="Italic">Pennatula phosphorea</Emphasis> (Anthozoa: Pennatulacea) from the west coast of Scotland

Reproduction in the sea pen Pennatula phosphorea Linnaeus 1758 was investigated in a population located in southern Loch Linnhe, west Scotland. This was accomplished through analysis of trends in oocyte size-frequency distribution and fecundity over a 12-month period. Pennatula phosphorea is dioecious and the study population exhibited a sex ratio of 1:1. Oogenesis in female P. phosphorea is characterised by the maintenance of a large pool of small oocytes throughout the year of which a small proportion (<30%) mature synchronously and are broadcast-spawned during the summer months of July and/or August. Although spawning occurs annually, the duration of oogenesis exceeds 12 months. Initial oocyte generation is best described as ‘quasi-continuous’ as it may be suppressed during winter. The timing and periodicity of oogenesis and spawning may be related to seasonal environmental cues and it is proposed that annual spawning constitutes a brief and synchronous event. Fecundity is high in P. phosphorea, typically up to 50 oocytes per polyp and 40,000 oocytes per colony in medium to large colonies. Although mean polyp fecundity increased with colony size (axial rod length) there was no seasonal fluctuation in this measure of relative fecundity. Since only a small proportion of oocytes present are spawned each year, the present study advises caution when making inferences regarding the seasonal output of viable oocytes, especially on occasions where the definition of fecundity or the details of the oogenic cycle of a species are unclear. Pennatula phosphorea produces large oocytes (>500 μm), indicative of the production of lecithotrophic larvae; this may confer certain advantages with respect to larval longevity and survival.     

Reproductive patterns in the Antarctic brachiopod Liothyrella uva

The reproductive cycle of the Antarctic articulate brachiopod Liothyrella uva (Broderip, 1833) is described from monthly samples collected between September 1985 and July 1987 from a population at Signy Island, Antarctica. Spermatogenesis and oogenesis are described for the first time in this species. Surface dried-tissue masses for a standard individual (41 mm shell length) were calculated for the digestive diverticula, gut, lophophore and gonad from monthly dissections of 15 brachiopods. Seasonal patterns, with summer peaks, were observed in the digestive diverticula and gut. The lophophore and gonad masses did not exhibit seasonal trends. Females showed a sharp decrease in proportion of large oocytes between October and November in 1986, suggesting spawning during this period. There was no similar decrease during the same period in 1985. Mean percent spermatozoa measurements revealed a large increase in November 1985 and a rapid decline in December 1985. This suggested a large spawning event for males in 1985 which was not repeated the following year. These data indicate large inter-annual differences in reproductive activity as well as differences between males and females. They also suggest the possibility of sperm storage by females. Brood characteristics were also highly variable. The smallest brooding female was 31.5 mm in length. Some females brooded more than one developmental stage simultaneously, and variation in brood size (numbers of embryos or larvae held in the lophophore) and brood composition between individuals was high. All samples collected throughout the 2 yr period contained some females with broods. The data suggest that the reproductive strategy of L. uva is highly plastic, and that there may be three reproductive periodicities on seasonal, annual and inter-annual time scales. Received: 16 May 1997 / Accepted: 2 March 1998     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). I. Gametogenesis and larval brooding

Anthelia glauca Lamarck, 1816 is a gonochoric, external-brooding soft coral found in KwaZulu-Natal. It is reproductively active in the summer months. The development of gametes produced in late summer is arrested in winter. Several stages of gametes are found at the base of the polyps, and female polyps produce several cycles of larvae over an extended breeding period of 4 to 5 months. Larvae are brooded in a unique pharyngeal brooding pouch not yet described in other coral species. The brood pouch consists of an expansion of the pharynx with constrictions proximal and distal to the embryos and larvae. Our data suggest that egg transfer and fertilization occur at full moon and the mature larvae are released after new moon. Zooxanthellae are absent in A. glauca oocytes, but zooxanthella infestation commences at the immature larval stage. Received: 15 July 1997 / Accepted: 12 March 1998     

Nutrition and development of brooded embryos in the brittlestar Amphipholis squamata: do endosymbiotic bacteria play a role?

To identify sources of nutrition potentially available to the yolk-deficient embryos of the brooding brittlestar Amphipholis squamata Della Chiaje, (= Axiognathus squamata, Thomas 1966), specimens were collected intertidally at all seasons (1986 through 1987) from Odiorne Point, Rye, New Hampshire and from Appledore Island, Isles of Shoals, Maine, USA. Ultrastructural, autoradiographic and immunohistochemical studies were made of adults and brooded embryos. Adult and embryonic tissues have morphological adaptations which support healthy, symbiotic bacteria (5 to 9 x 10⁶ colony forming units/adult brittlestar), while autoradiographic studies indicate direct uptake of labeled amino acids by eukaryotic host tissues and bacteria. Cell envelopes of subcuticular bacteria suggest that they are Gram negative and may belong in the genus Vibrio. Based on immunohistochemical localization, it appears that a single type of bacterium is present in large numbers under the cuticles of embryos and adults of A. squamata. This is the first study of the relationship between an echinoderm and a bacterium which includes isolation and immunohistochemical verification of the identity of the bacterial symbiont.     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

Embryogenesis and acquisition of algal symbionts by planulae of Xenia umbellata (Octocorallia: Alcyonacea)

Early embryogenesis of the internally brooding soft coral Xenia umbellata and acquisition of algal symbionts in the course of its planular ontogenesis have been examined by scanning and transmission electron microscopy and by light microscopy. The endoderm of adult X. umbellata harbours symbionts mainly in the tentacles and in the peripheral solenia system. The colonies are gonochoric brooders. Algal symbionts were never found in the sperm sacs, and were only rarely found in the follicular tissue enclosing the oocytes. Fertilized eggs pass into endodermal brood pouches where embryogenesis occurs. Cleavage is holoblastic and leads to formation of a solid blastula. Algal symbionts are conspicuously embedded in the parental mesoglea that coats the young embryo, most probably transmitted by surface adherence. At a further stage, this integument disappears and the algae reside extracellularly among the cells of the newly-formed blastula. After subsequent cell proliferation developing planulae possess an inner mass of yolk-laden cells that contain numerous symbiotic algae. Gradually the yolk disintegrates, leaving a cavity enclosed by ectoderm, a thin mesoglea and an inner endoderm with intracellular symbionts. The mature planulae have already been provided with numerous intracellular symbionts by the time they are expelled from the brood pouches. The markedly early symbiont acquisition by the embryos of X. umbellata may help support their developmental requirements in the course of planular ontogenesis.     

The reproductive biology of Hymenaster membranaceus from the Rockall Trough,North-East Atlantic Ocean,with notes on H. gennaeus

Samples of the deep-sea spinulosan asteroid Hymenaster membranaceus Wyville Thomson were collected in a timeseries of 19 bottom trawls spanning the period April 1978 to October 1981 from a 2200 m-deep station in the northern Rockall Trough. The reproductive biology of this species was studied from histological sections of the gonad, and compared with that of H. gennaeus H. L. Clark which was collected in the same hauls. At first sexual development, oogonia develop in nests surrounded by small accessory cells. Previtellogenic oocytes remain in the periphery but, at maturity, oocytes ranging up to 1 100 m fill the ovary. A variety of accessory cells pack the lumen and may be nutritive or degenerative. It appears that a small number of oocytes are spawned intermittently, but there is no evidence of overproduction and break-down of superfluous oocytes. A few large oocytes become senescent and undergo internal break-down, releasing periodic acid Schiff-positive material into the lumen. Size-frequencies of oocytes indicate that eggs may be spawned as a continuous slow release, and there is no evidence of reproductive synchrony between or within samples. On reaching maturity, males appear always to be ready to release spermatozoa. Spawning is probably stimulated by egg release during chance encounters with mature females. There is no evidence for brooding, and from the large size and yolky nature of the egg direct lecithotrophic development at or near the seabed is inferred. A limited histological study of H. gennaeus indicates that egg production is very similar, but the two species differ in the nature of the accessory cells and amorphous material filling the lumen.     

Reproductive ecology of the brooding,hermaphroditic clam Lasaea subviridis

The reproductive ecology of a small intertidal clam was studied for two years (September 1980-November 1982) at two mid-intertidal sites on the northern shore of Monterey Bay, California. Lasaea subviridis is hermaphroditic and incubatory; the young are released as juveniles. Brooding adults are found throughout the year. All embryos in a clam are at the same stage of development, but the number and stage of embryos varies among clams at any one time. Individuals are iteroparous and produce small numbers of oocytes and sperms simultaneously. All stages of gonad development are found in different individuals at any one time. There was no evidence of seasonality. Young reared in the laboratory after being dissected from the parent reached brooding size (shell length of 1.7 mm) within 6 mo. Laboratory experiments indicated that juveniles crawl away soon after release from the parent. These reproductive characteristics may contribute to this taxon's ability to colonize new environments rapidly.     

Demography and reproductive biology ofMelinna palmata (Ampharetidae: Polychaeta) in Inner Galway Bay on the west coast of Ireland

A high-density population of the polychaeteMelinna palmata Grube was the focus of a two-year study at a shallow-water location in Inner Galway Bay on the west coast of Ireland. Mean densities were highest in the second year of the study following successful recruitment. Size-frequency analysis of population structure (April 1983 to March 1985) revealed the presence of four cohorts, with the 1983 settlement becoming the dominant modal class. Size comparison of this cohort with the 1982 settlement showed that growth rates were lower and mortality higher in the 1983 settlement, which may be related to density dependent intra-specific competition.M. palmata at this location live for 2 to 2.5 yr with a small number surviving to 3 yr of age.M. palmata in Galway Bay is dioecious and has an even sex ratio. Reproductively mature worms were approximately 2 yr old when they first spawned. Spawning took place between May and July, when mature oocytes had a modal size of ca. 290µm. Proliferation of previtellogenic oocytes (ca. 10µm) from the gonads occurred following spawning. Growth over the winter was slow and was followed by a period of rapid growth between March and May, coincident with rising water temperatures. The presence of mature unspawned oocytes, which were resorbed after commencement of new proliferation, was taken as evidence of the species propensity for polytely. However, post-spawning mortality appeared to be high, so that only a small number of individuals survived to spawn the following year. The highest abundance of mature males was encountered between May and August, but lower levels of mature stages were present throughout the year.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

玉米赤霉烯酮暴露对秀丽线虫生殖系统的损伤作用

环境雌激素玉米赤霉烯酮(ZEN)的生态安全性近年来受到越来越多的关注。以秀丽隐杆线虫(Caenorhabdites elegans)为模式生物研究ZEN对线虫的生殖损伤作用。在DIC显微镜下观察,48 h暴露后与对照组相比,暴露组线虫性腺臂明显萎缩,卵母细胞数目和大小均受到显著影响(P0.01),并且可见部分暴露组线虫产卵器严重畸形。这均表明ZEN暴露对秀丽隐杆线虫生殖系统的发育和功能可产生损害作用。     

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.     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent     

Reproductive habits and life cycle of the small clam Kingiella chilenica (Bivalvia: Cyamiidae) in an estuarine sand flat from the south of Chile

A population of the small clam Kingiella chilenica Soot Ryen 1959 was studied from March 1986 to December 1988 in an intertidal flat at Queule River, in the south of Chile. The life cycle and life history pattern of the bivalve were established contrasting population structure and dynamics to its reproductive habits. Individuals are gonochoristic and semelparous, presenting a typical annual life cycle. The species is a sequential brooder whose embryos undergo direct development. After the brooding season (summer through autumn), the adults disappear gradually (autumn through winter). Recruited juveniles overwinter during a relatively long period, undergoing rapid growth during the spring to attain the adult stage during the summer. The number of brooded embryos increases in proportion to adult length cubed. Life history traits of this bivalve are compared to those reported for other small brooder clams. Some basic tendencies become apparent when traits for semelparous versus iteroparous species are contrasted. As in other semelparous sequential brooders (Transennella tantilla, Gaimardia bahamondei), the relationship between brood size and shell length obseved in K. chilenica does not fit the allometry hypothesis for marine brooding invertebrtes that allometric constraints on the brooding space limit the fecundity of larger individuals. Also contrary to theoretical predictions, small body size does not limit the diversity of a clam's developmental patterns. Possible explanations for this finding are discussed.     

Slow developing demersal embryos and larvae of the antarctic sea star Odontaster validus

The early development of Odontaster validus at McMurdo Sound, Antarctica, is indirect and includes equal cleavage, a convoluted blastula, a free-swimming coeloblastula, a gastrula, and a feeding bipinnaria larva. Development differs from that of other asteroids in two respects: (1) The developmental rate is extremely slow; blastulae form nearly 2 days after fertilization, gastrulation begins after 7 days, and the bipinnaria develops in about 40 to 55 days. The slow developmental rate appears to be only partly related to the low environmental temperature (-1.5°C). (2) The embryos and larvae are largely demersal. Such behavior may be an adaptation to keep the larvae out of antarctic surface waters, as does brooding in many other polar echinoderms.