Effects of temperature,nutritive and metal stressors on the reproductive biology of Mytilus edulis

Experimental observations suggest that the gametogenic cycle of Mytilus edulis from Swansea Bay, South Wales (U.K.) is adversely modified by the combined effects of temperature, nutritive and sublethal metal-induced stress. During low temperature stress, oogenesis remained in the vegetative stage and vitellogenesis only commenced when temperatures were raised to ambient levels. Copper was the most toxic metal, although its uptake was slowest. A concentration of 0.05 mg l^-1 of this metal proved lethal at ambient temperatures, although at low temperatures there was no mortality. Copper suppressed both the growth of young oocytes and vitellogenesis in larger oocytes. Zinc was less toxic than copper, although this also inhibited oocyte development and resulted in severe lysis of gametes. Cadmium was the least toxic of the three metals studied and suppressed gametogenesis only in the initial stages of gonad development. The low toxicity of cadmium to gametogenesis may be a result of its high solubility and its possible storage in the digestive gland.     

Reproductive cycle of the endemic cassiduloid<Emphasis Type="Italic"> Cassidulus mitis</Emphasis> (Echinoidea: Cassiduloida) on the Brazilian coast

Cassidulus mitis Krau, 1954 is an endemic species from Rio de Janeiro, Brazil. To study the reproductive cycle of this species, samples were collected at Praia Vermelha, in Guanabara Bay, from October 1998 to April 2000. The sex ratio for C. mitis was 1:1, and reproduction occurred throughout the year. The gonadal index of males was greater than that of females. Five gametogenic stages were identified in males: early growth, premature, mature, partially spawned and early growth with a partially spawned stage. There were no spent or recovery stages such as those seen in females. Females had six gametogenic stages: recovery, early growth, premature, mature, partially spawned and spent stage; there was no early growth with a partially spawned stage as there was in males. The mean oocyte diameter was 382 m (SD=49 m), and the mean number of juveniles per female was 100 individuals (SD=108). The anterior gonads were frequently smaller than the posterior ones, and there were differences in the gametogenic stages between the anterior and posterior gonads. First sexual maturation occurred in individuals with a diameter of 17 mm. The intestinal wet weight was two- to threefold higher than the test wet weight. This may indicate that C. mitis uses its intestine as a weight belt to avoid displacement from the substratum.Communicated by P.W. Sammarco, Chauvin     

Reproductive ecology of <Emphasis Type="Italic">Fungiacyathus marenzelleri</Emphasis> from 4100 m depth in the northeast Pacific Ocean

Fungiacyathus marenzelleri (Vaughan, 1906) is a deep-water solitary coral, cosmopolitan in distribution that is found at depths of 300–6,328 m. This study examined gametogenesis, inter-annual variability and reproductive periodicity of F. marenzelleri collected from Station M (34°50′N, 123°00′W) in the northeast Pacific at a depth of 4,100 m. Samples were collected (May, June, October 1996; August 1998; February, June 2001; and June 2002) and histologically processed with spermatogenic stage, oocyte size and fecundity measured. Four stages of spermatogenesis were identified and all males contained multiple stages of sperm development in each seasonal sample. Three stages of oocyte development were identified; previtellogenic (<28–150 μm), vitellogenic (150–300 μm) and late vitellogenic (300–400 μm). Comparison of mean oocyte diameters among sampling dates showed there were no inter-annual variations or seasonal differences. Overall, fecundity was 1,290 (±407) oocytes polyp⁻¹, and with no significant differences between sample months. Fecundity was not polyp-size dependent. This study shows a similar quasi-continuous mode of reproduction to this species examined from the Northeast Atlantic Ocean, but the fecundity is reduced by 50%. The reproductive output may fluctuate in relation to the input of organic material at this site, as shown by non-significant trends in the oocyte size-frequency and fecundity data. A quasi-continuous output of gametes would promote successful fertilisation and wide dispersal of the lecithotrophic larvae.     

Influence of habitat on the reproductive biology of the deep-sea hydrothermal vent limpet <Emphasis Type="Italic">Lepetodrilus fucensis</Emphasis> (Vetigastropoda: Mollusca) from the Northeast Pacific

Habitat selection by the hydrothermal vent limpet, Lepetodrilus fucensis, in Northeast Pacific hydrothermal vent ecosystems, may influence its reproductive output, as it occupies habitats with varying physico-chemical conditions that reflect the availability of nutritional resources. Histological techniques were used to determine size at first reproduction, gametogenesis, reproductive output, and fecundity in relation to shell length (SL), through examination of the gonads of male and female L. fucensis, collected from five different hydrothermal vent habitat types with different temperature anomalies and hydrothermal fluid flow vigour: vigorous (VIG), diffuse (DIF), tubeworm bushes (TWB), peripheral (PER), and senescent areas (SEN). Both male and female L. fucensis exhibited early maturity, with the first reproductive event occurring at 3.8 and 3.9 mm shell length, respectively. All stages of gamete development were present in the gonads of males and females, suggesting continuous gametogenesis and asynchronous reproduction in this species. Gametogenic maturity of limpets did not vary among actively venting habitats (VIG, DIF, TWB, and PER), but was significantly lower in males and females from SEN habitats. Mean oocyte diameter was largest in females from VIG habitats, and smallest in females from SEN habitats, than in those from the other habitats (DIF, TWB, and PER). Females from actively venting habitats also had greater actual fecundity than those from senescent habitats. While the gametogenic pattern of L. fucensis appears phylogenetically constrained, selection of actively venting habitats by L. fucensis maximizes its reproductive output. The multiple feeding strategies of L. fucensis may allow for a constant supply of energy to be allocated to reproduction in any habitat except senescent vents. Early maturity, high fecundity, and continuous production of gametes suggest a reproductive strategy characteristic of an opportunistic species, and may be contributing to the extremely abundant populations of L. fucensis observed in the Northeast Pacific vent ecosystem.     

Introduction of the northern Pacific asteroid Asterias amurensis to Tasmania: reproduction and current distribution

The northern Pacific asteroid Asterias amurensis (Lütken) has been recently introduced to Tasmania and is now well-established in east and southeast Tasmania. This sea star is conspicuous throughout the Derwent River estuary and is particularly abundant in the Hobart port area. Reproduction of two populations of A. amurensis at Sullivans Cove and at Sandy Bay was investigated from August 1993 to October 1994 by gonadal histology and measurement of the gonad index (GI). An intense period of vitellogenic and spermatogenic growth started in April, with the breeding condition reached by June. In 1994, peak GI was recorded at Sullivans Cove in July and at Sandy Bay in August. Although these data indicated that the Sullivans Cove population spawned before the Sandy Bay population, histological examination revealed that major spawning activity occurred in both populations from July onwards. In the early part of the breeding season, gametogenesis proceeded in parallel with spawning, with released gametes being replaced by continual gametogenesis. Maintenance of a higher GI during the early part of the breeding season at Sandy Bay was due to prolonged gametogenic replacement at this site. Spawning continued to October 1994, resulting in a sharp decline in the GI. The cyclic expansion and regression of the genital haemal sinus coincident with germinal proliferation and growth, respectively, support the contention that haemal fluid provides nutrients for gametogenesis. In Tasmania, A. amurensis experiences temperature and photoperiod regimes similar to those experienced by endemic populations of this species in the north Pacific. Comparison of reproduction of A. amurensis in Japan and Tasmania at similar latitudes shows that gametogenesis of the northern and southern populations is 6 mo out of phase. This phase shift provides evidence for photoperiodic regulation of gametogenesis in A. amurensis. The similar trend in sea-temperature regimes associated with the stages of gametogenesis in northern and southern populations suggests that temperature also plays a modulatory role. Based on the periodicity of reproduction and the ontogeny of A. amurensis, the planktotrophic larvae of this species would be expected to be in the plankton for several months from the end of winter through summer. Considering the highly fecund, dispersive life-history of A. amurensis, the southeast Tasmanian populations have considerable potential to serve as a seed source for establishment of new populations of this asteroid elsewhere in Tasmania and in mainland Australia. Received: 18 October 1996 / Accepted: 22 October 1996     

Can photoperiod manipulation affect gonad development of a boreo-arctic echinoid (Strongylocentrotus droebachiensis) following exposure in the wild after the autumnal equinox?

A laboratory experiment was conducted in the winter of 2003–2004 to assess the effect of varying photoperiod regime on consumption rate, assimilation rate, absorption efficiency, and gonad development of the green sea urchin, Strongylocentrotus droebachiensis. Adult individuals were collected from the wild after they had been exposed to the ambient autumn photoperiod cue (which is the extraneous trigger thought to elicit gametogenesis in this species) and placed at ambient temperature for 12 weeks under five different photoperiod regimes: (1) 24 h light:0 h dark=“0D”, (2) 16 h light:8 h dark=“8D”, (3) 8 h light:16 h dark=“16D”, (4) 0 h light:24 h dark=“24D”, and (5) ambient photoperiod (range: 10.50–15.25 h dark). Urchins in these five treatments were fed ad libitum with bull kelp, Nereocystis luetkeana. A sixth treatment consisted of starved individuals held under 0D conditions. Various gonad factors including gonad index, percent gonad water, gonad colour (CIE lightness or L*, CIE hue or a*, and CIE chroma or b*), percent area occupation of the gonad by various cell types (nutritive phagocytes, spermatozoa, and secondary oocytes/ova), and stage of development were assessed at the beginning of the experiment and at weeks 4, 8, and 12 of the study. Consumption and assimilation rates were assessed at weeks 4 and 12 and absorption efficiency at week 12 of the experiment. Urchins were predominantly in the growing and premature stages at the beginning of the experiment, but by week 4 at least 20% of individuals in all treatments receiving food were classified as mature. Spawning occurred during all these treatments between weeks 4 and 8, as evidenced by significant decreases in spermatozoa and secondary oocytes/ova and a significant decrease in percent gonad water, but was not accompanied by major declines in gonad indices. Greater than 90% of individuals in all five of the fed treatments were in the recovering and growing stages at the end of the experiment. The 16D treatment had by far the greatest percentage of urchins in the growing stage. In contrast, individuals that were starved were predominantly in the mature stage at weeks 4, 8, and 12, with only ~30% reaching the spawning stage by the end of the experiment. Photoperiod significantly affected gonad indices at the termination of the experiment with gonad index being the highest in the 16D treatment; this was significantly greater than in the 8D and ambient treatments. Photoperiod did not significantly affect gonad percent water, gonad lightness, or gonad hue. Gonad chroma was significantly affected by photoperiod, urchins held under ambient conditions having significantly lower b* readings than individuals in any other treatment. Photoperiod had little or no affect on consumption rate, assimilation rate, or absorption efficiency. Thus, differences among treatments in regards to gonad index, gonad chroma, and stage of development cannot be attributed to variations in feeding, absorption, or assimilation. The results of this experiment indicate that once gametogenesis is initiated, photoperiod manipulation cannot prevent ultimate spawning. However, photoperiod regime can affect the rate at which urchins move through the various stages of the gametogenic cycle. Urchins placed on short days under artificial lighting (16D) moved through the spawning stage into recovering and growing stages the fastest. This photoperiod regime also produced the highest gonad index at the end of the experiment. Since the commercial urchin market prefers large gonads in the growing and premature stages (i.e. before the mature stage is reached and gonads start leaking sperm and eggs), short day-lengths under artificial lighting (16D) appear to be the best photoperiod conditions for optimizing marketability.     

Gametogenesis of Pollicipes cornucopia (Cirripedia: Scalpellomorpha) in north-west Spain

The gametogenesis and gonad microanatomy of the stalked barnacle Pollicipes cornucopia (Leach) is described. An histological classification of gametogenic development is proposed for the firt time for a species of the Cirripedia. This species has an annual gametogenic cycle on the north-west Spanish coast. Spawning takes place mainly from March to September. Ovaries release gametes at least twice during the gametogenic cycle.     

Reproductive periodicity in the euryalinid snake star Astrobrachion constrictum in a New Zealand fiord

The euryalinid snake star Astrobrachion constrictum (Farquhar), a normally deep-water species, exists within SCUBA diving range in New Zealand flords, and thus presents a unique opportunity for study. A population of A. constrictum in Doubtful Sound was investigated from February 1990 to July 1994 to determine the reproductive cycle. Unlike most ophiuroids, the gonads of A. constrictum are positioned along the aboral coelom in the proximal portion of the arms. The proximal 2 to 3 cm of the arms of individuals were thus removed on an approximately monthly basis for histological examination of the gonads. Image analysis was used to determine oocyte size-frequency distributions. A gonad index was calculated for each individual based on the ratio of cross-sectional area of the gonads to cross-sectional area of the arm. Maturity indices were calculated based on stage of development of the gonads of each individual. A. constrictum has a period of enhanced breeding activity in April–June, after which both the gonad and maturity indices decline. The increase in gametogenic activity during these months was correlated with rising sea temperature within the fiord. Although the females showed an annual pattern of gonad development, oogenesis was continuous throughout the year, with oocytes at different stages of development present in the gonads. Males also showed continuous spermatogenesis and, although the same annual pattern of gonad development was evident as seen in the females, it was less marked. The presence of females with large oocytes and ripe males throughout the study indicates that breeding in the Doubtful Sound population is prolonged, and possibly continuous, as found in many deep-sea ophiuroids.     

When fragmented coral spawn? Effect of size and timing on survivorship and fecundity of fragmentation in <Emphasis Type="Italic">Acropora formosa</Emphasis>

In order to determine competency of sexual reproduction and survival rate after fragmentation, the branching coral Acropora formosa was fragmented and fragments in three different sizes (ca. 5, 10 and 20 cm long) and three different stages of gametogenesis were transplanted on coral pavement. Their oocyte development and fecundity, as well as spawning were monitored for a 3-year period. The oocyte development was affected by both fragment size and by the developmental stage of oocytes when fragmented. In small fragments, the oocytes were resorbed while in large fragments they continued development. Oocytes in the early vitellogenic stage at the time of fragmentation were resorbed, whereas those in the late stage continued developing. Smaller fragments showed a lower survival rate and histological observations of their gonads revealed resorption of oocytes, suggesting that there was a trade-off of energy between reproduction and survival. Transplanted fragments often spawned one month earlier than the donor colonies in the first year, but spawning occurred in the same month as the donors or did not occur at all in the second year and none spawned in the third year. The risk of colony death may cause the fragments to re-allocate energy for sexual reproduction.     

The reproductive biology of echinothuriid and cidarid sea urchins from the deep sea (Rockall Trough,North-East Atlantic Ocean)

The reproductive biology of 5 species of echinothuriid (Phormosoma placenta, Calveriosoma hystrix, Araeosoma fenestrum, Sperosoma grimaldii and Hygrosoma petersii) and 2 species of cidarid (Cidaris cidaris and Poriocidaris purpurata) sea urchins from the deep sea (Rockall Trough) has been examined from samples collected during 1973–1983. In all species the gonads lie within the interambulacrum attached to aboral gonopores and when fully developed occupy most of the test not occupied by the gut or Aristotle's lantern. In all the species, initial oocyte development takes place along the germinal epithelium embedded in nutritive tissue. In all the echinothuriids and in Poriocidaris purpurata, the oocyte grows to ca. 200 to 450 m, at which stage vitellogenesis begins. Oocyte growth continues until a maximum egg size of 1 100 to 1 500 m is attained. In the echinothuriids, two types of nutritive tissue are found. In the carly stages of gametogenesis the oocyte is surrounded by well-structured periodic acid Schiff (PAS)-positive tissue. As the oocyte grows this tissue becomes vacuolated, suggesting that there is a transfer of nutriment to the developing oocyte. In Phormosoma placenta, unspawned oocytes are phagocytosed. There is no evidence of seasonality in any of the echinothuriid species or in Poriocidaris purpurata. Extrapolation with shallow-water echinothuriids suggests that larval development is lecithotrophic, omitting any planktotrophic phase. Of the species examined, only Cidaris cidaris has a reproductive strategy which produces a known larva, although the limited samples did not permit any determination of seasonality in this deep-sea population.     

Reproductive cycle and gametogenesis in the black abalone Haliotis cracheroidii (Gastropoda: Prosobranchiata)

Seasonal changes in gonald size and stages of gametogenesis in the black abalone Haliotis cracheroidii were related to changes in environmental parameters. H. cracheroidii showed an annual reproductive cycle terminating in a synchronized spawning in late summer. Gametogenesis was initiated immediately after spawning. Gametes were present in the gonad through the winter months. Gametogenesis was initiated a second time in the spring months. Maximal gonad growth (to a gonad index of 20%) occurred during summer months prior to spawning. Changes in gonad size andperiods of initiation of gametogenesis revealed no apparent correlation with changes in seasonal water temperature. Changes in gonad size showed no apparent relation to change in day length. Total polysaccharide levels in foot tissue changed seasonally, indicating that food availability is probably not a factor in directly regulating gonad growth. Gonad index data for the chiton Katharina tunicata (collected over a 10 year period) showed no apparent correlation to seasonal change in water temperature.     

Spatio-temporal population and reproductive responses in Perinereis gualpensis (Polychaeta: Nereididae) from estuaries under different anthropogenic influences

Reproductive and population-level responses are ecologically relevant diagnostic tools for revealing the cost of long-term tolerance to contaminants. Spatial density distribution, seasonal density, biomass, size frequency histograms and oocytes stages in Perinereis gualpensis (Polychaeta: Nereididae) from a highly anthropogenically impacted estuary (Lenga, south-central Chile) were compared with low-polluted estuaries (Tubul and Raqui). Spatial distribution showed that the highest P. gualpensis abundances occur in the central (middle) estuarine area, establishing them as suitable zones for comparisons of estuaries. Middle areas also showed differences among estuaries in terms of population and reproductive responses. Mature stage oocytes and juvenile recruitment limited to certain seasons and low adult survival led to significant low densities and biomasses in Lenga individuals (p?相似文献   

Reproduction in five species of crinoids at Lizard Island,Great Barrier Reef

Reproduction in five species of crinoids [Himerometra bartschi (A. H. Clark), H. robustipinna (P. H. Carpenter), Cenometra bella (Hartlaub), Colobometra perspinosa (P. H. Carpenter), and Oligometra serripinna (P. H. Carpenter)] was studied at Lizard Island, Queensland, over 24 mo, from April 1981 to March 1983 (except for O. serripinna —June 1981 to March 1983). All five species of crinoids were dioecious, although a few instances of synchronous hermaphroditism were noted in H. bartschi, C. perspinosa, and O. serripinna. Gonad morphology and the sequence of developmental stages were similar in each species. Within individuals, gametogenesis was synchronous in gonads both along and between arms, except in some proximal and distal genital pinnules in which gonads never matured. Within a species, there was a degree of synchrony of gametogenesis in a sample, but the level of gametogenic activity varied considerably between years. A high level of continuous reproductive activity was observed in H. bartschi, H. robustipinna, Cenometra bella, and Colobometra perspinosa from about mid-summer to early/mid-winter, while O. serripinna probably had a bimodal reproductive cycle with peaks around February and June. There was a significant 1:1 male:female sex ratio in the populations sampled, and unsexable individuals were significantly smaller (as determined by maximum arm length) than either sexable males or females.     

Reproductive biology of the commercial sea cucumber Holothuria fuscogilva in the Solomon Islands

 Reproduction of Holothuria fuscogilva (Selenka, 1867) in the Solomon Islands was investigated over a 4 yr period (1994 to 1998) by macroscopic and microscopic examination of the gonad tubules, the gonad index (GI) method, histological examination of gametogenesis, and spawning-induction trials. The gonad consisted of numerous tubules that dominated the coelom of gravid specimens. New tubules appeared in March, and grew in size and extent of branching until they reached their maximum size and maturity in August. Spawning occurred from August to October, with the majority of gametes released during October, although it was only partial in many individuals. After spawning, the tubules appeared wrinkled and resorbed into the gonad basis. A five-stage gonad maturity scale based on the macroscopic appearance of the gonad tubules corresponded with discrete stages of gametogenesis identified by histology. Gametogenesis was initiated in mid-March, with oogenesis and spermatogenesis occurring in parallel, followed by the growing stage (May to July) which was marked by active gamete development. Successful induction of spawning during the breeding period corroborated the GI and histological data. The uniform growth of gonad tubules indicated that H. fuscogilva in the Solomon Islands does not conform to the progressive tubule recruitment model described for other holothurians. An important application of this study is that the appearance of gonad tubules, removed by biopsy, can be used to determine the gonad condition of wild adults or captive broodstock. Received: 28 June 1999 / Accepted: 8 March 2000     

Ovarian allometry and the use of the gonosomatic index: a case study in the Mediterranean sardine,<Emphasis Type="Italic"> Sardina pilchardus</Emphasis>

The gonosomatic or gonadosomatic index [GSI=(ovary weight/fish weight)×100] has been widely used in fisheries science and experimental reproductive studies as a simple, low-cost measure of reproductive condition. However, its properties have not been fully evaluated, and several pitfalls, such as size-dependence and changes in ovarian allometry, may invalidate its use. In the present study, we examined ovarian allometry and the appropriateness of GSI for assessing ovarian activity in the Mediterranean sardine Sardina pilchardus. The analysis was based on a large sample of histologically scored females collected over an annual cycle in the eastern Mediterranean (Aegean and Ionian Seas). First, we examined GSI dependence on fish size by comparing ovary weight–on–fish weight relationships in different stages of oocyte development. The effects of recent spawning (incidence of postovulatory follicles) and intensity of follicular atresia were also addressed. In a subsequent step of the analysis, we applied generalized linear models (GLMs) to analyze the relationship between GSI and histological stage, taking into account the parallel effects of additional factors related to geographical region, month of capture, fish length and energetic reserves (fat stage, hepatosomatic index—HSI). Ovarian growth was isometric in all stages of oocyte development and states of atresia, but altered to positive allometric at the stage of oocyte hydration. Oocyte growth and intensity of atresia significantly affected GSI. Fish length, geographical region, month of capture, fat stage and HSI did not substantially affect GSI, further strengthening the appropriateness of the index for the Mediterranean sardine. Finally, we provide first evidence, based mainly on available (but yet limited) published information for other fish species, that: (1) the pattern of ovarian allometry may not be altered by fecundity variations and (2) the pattern of ovarian allometry may reflect the pattern of oocyte growth (i.e. isometric vs. allometric ovarian growth could reflect group-synchronous vs. -asynchronous oocyte development). Alternatively, the pattern of ovarian allometry could reflect the presence or absence of size dependency in the relative fecundity (eggs per gram of body weight) and/or egg size. The isometric ovarian growth in the Mediterranean sardine could be attributed to the lack of such size-dependent effects.Communicated by O. Kinne, Oldendorf/Luhe     

Annual reproductive cycle of the green sea urchin, Strongylocentrotus droebachiensis, in differing habitats in Nova Scotia, Canada

We monitored the reproductive cycle of Strongylocentrotus droebachiensis (OF Müller) between April 1993 and August 1995 in kelp beds, barren grounds and grazing fronts at both a wave-exposed and a sheltered site along the Atlantic coast of Nova Scotia. Gonad index and histological analyses showed that S. droebachiensis has an annual reproductive cycle that is synchronous across sites and habitats, and between females and males. Spawning occurs in March/April of each year but a small proportion of sea urchins in the study populations also spawned in fall 1995. During most of the year, sea urchins in kelp beds and grazing fronts have a higher gonad index than those in barren grounds. Gonad indices also tended to be higher at the wave-exposed than the sheltered site. Interannual variability in peak gonad index was significant in the barren grounds at the wave-exposed site and in the grazing front at the sheltered site. The gametogenic cycle is characterized by six stages based on the abundance of nutritive and germinal/gametic cells. Nutritive phagocytes are abundant after spawning and replaced by increasing numbers of germinal and gametic cells as the gametogenic cycle progresses. The temporal patterns of abundance of each cell type were similar among habitats indicating that the gonads were qualitatively similar despite large differences in gonadal mass. The quantity of gut contents (ratio of food volume to body volume) was similar among habitats, but the quality (percentage of organic material) tended to be higher in kelp beds and grazing fronts than in barren grounds suggesting that differences in gonad index of S. droebachiensis in different habitats are related to differences in diet. The high density of sea urchins in grazing fronts combined with their high fecundity suggests that they make the greatest contribution, per unit area, to the overall larval pool. Received: 20 May 1997 / Accepted: 21 January 1998     

The annual reproductive cycle of the holothurian Aslia lefevrei (Dendrochirota: Echinodermata)

The dendrochirote holothurian Aslia lefevrei (Barrois) is locally and densely aggregated (>70 individuals m^-2) in Galway Bay, on the west coast of Ireland. It has a discrete, relatively short breeding period from the end of February to the beginning of April, with peak activity in March. The sexes are separate and the single gonad, consisting of numerous tubules (100 to 800), lies in the anterior coelom. Both gonad index values and histological observations indicate that gametogenesis resumes in June/July after a brief post-spawned resting period. Little oocyte growth and sperm production occurs over the warm summer months when the holothurian is actively feeding. Gonad index values, however, increase rapidly during this period and reach high values by early winter without concomitant increases in maturity indices. This is subsequently followed by slight reductions in the gonad indices, before reaching peak values in February. It is suggested that this is probably due to the initial build-up and subsequent assimilation of storage products in the gonad, the development of gametogenic products and the frequent assimilation thereafter of unwanted ova. Spawning generally takes between 5 and 60 min, with the entire contents of the gonad rarely being shed in a single release. During this process, the holothurian raises its anterior end and, with the extended tentacles close together, emits a stream of gametes from the genital pore in the mid-dorsal line. Laboratory experiments and general observations indicate that increased temperature and low light levels may influence spawning behaviour.Contribution No. 248 to the School of Marine Sciences, University College Galway, Ireland     

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     

Oocyte development in the kuruma prawn Penaeus japonicus

Female kuruma prawns (Penaeus japonicus Bate) with undeveloped, early developing, developing, nearly ripe and ripe ovaries, were collected from Ise Bay, Japan, in 1984. Oocyte development of the kuruma prawn was classified into ten stages according to morphological characters, namely: (1) synapsis stage, (2) chromatin nucleolus stage, (3) early perinucleolus stage, (4) late perinucleolus stage, (5) oil globule Stage I, (6) oil globule Stage II, (7) yolkless stage, (8) yolk granule stage, (9) prematuration stage, and (10) maturation stage. The synapsis stage is a multiplication stage. The chromatin nucleolus stage, early and late perinucleolus stages are previtellogenesis and primary growth stages. Oil globule Stage I is an initial stage of primary vitellogenesis and secondary growth. Follicle cells on the oil globule Stage I oocytes expand rapidly and reach maximum size during oogenesis. Yolk granule stage oocytes are in the initial stages of secondary vitellogenesis. Strongly acidophilic yolk granules accumulate within basophilic vesicles of the cytoplasm. The yolk granules are first concentrated in the inner part of the cytoplasm, then gradually spread to the periphery. Cortical crypts, which are separated from the oocyte cytoplasm by the cytoplasmic membrane, are situated outside of oocyte cytoplasm. Germinal vesicle breakdown (GVBD) is initiated in the late phase of prematuration and continues until the late phase of maturation immediately prior to spawning. At the beginning of the maturation stage, the oocytes are ovulated, after which the nuclei further shrink and migrate out-wards. After ovulation, meiotic division of the ovarian oocyte progressed up to the metaphase of primary maturation division. Finally, the meiotic metaphase is visible just beneath the cytoplasmic membrane in the mature oocyte. Though ovulation is synchronous within the same ovary, GVBD is not completely synchronous. Ovulated mature oocytes have many club-shaped cortical crypts in the peripheral part of the cytoplasm and contain extensive accumulations of yolk granules dispersed throughout the cytoplasm. The apical end of the club-shaped cortical crypts and cytoplasmic membrane are coated by the vitellin envelope in the mature oocyte.