Long-term changes in populations of subtidal bivalves (Abra alba and A. prismatica) from the Bay of Morlaix (Western English Channel)

Two populations of Abra alba (Wood) and one of A. prismatica (Montagu) (Mollusca: Bivalvia) were studied over a 10 yr period (1977–1987) in two muddy fine-sand subtidal communities of the Bay of Morlaix, France. The survey provided an example of long-term changes in the three Abra spp. populations, which displayed synchronized changes, with a regular annual cycle and increasing densities during 1979–1980 related to the higher concentration of organic matter resulting from the Amoco Cadiz oil spill in March 1978. A. alba rapidly adapts its demographic strategy to eutrophic conditions by increasing its reproductive potential, growth, and abundance. During times of eutrophication, A. alba has three spawning periods and three recruitments per year as opposed to two spawning periods and two recruitments per year during oligotrophic conditions. Growth of the juveniles of this species is insignificant until spring for individuals recruited in the autumn, whereas individuals which settle during spring or summer display immediate rapid growth. A. prismatica has a low capacity to adapt to eutrophic conditions. It has one annual period of sexual maturation at the end of the summer, with spawning in September–October and settlement beginning in mid-November. Growth of the juveniles after settlement is also insignificant until April. These results enable comparison of the demographic strategies of these two sympatric species.     

Growth patterns of Amphiura filiformis support the hypothesis of organic enrichment in the Skagerrak-Kattegat area

Population size structure, abundance and biomass of the burrowing ophiuroid Amphiura filiformis were measured in the period 1970 to 1989 at three widely spaced sites at 55 to 100 m depth in the Skagerrak-Kattegat area. This species is a dominant component of the fauna in the area, and also one of the species accounting for a great part of a recent general increase in benthic biomass. Abundance at two of the sites and biomass at all three sites increased significantly with time from the 1970s onwards. Maximum individual size and growth of the cohort of adults showed a marked increase in the period 1980 to 1983 at the three sites. In the 1980s annual absolute growth increments of the recruits were clearly greater than in the 1970s, and size-specific growth rates were virtually unchanged. These results are consistent with the hypothesis that recent increases in benthic biomass in this area result from increased input of food to the benthos. A. filiformis has the potential for rapid growth; estimated maximum weight-specific growth rates were more than 1.5% d^-1 in the disc diameter interval 3 to 9 mm, which is comparable to rates found in shallow water filtering bivalves. The maximum disc diameter growth rate was ca. 0.4 mm mo^-1.     

Reproductive cycle of<Emphasis Type="Italic"> Bispira volutacornis</Emphasis> (Polychaeta: Sabellidae) on the west coast of Ireland

The reproductive cycle of Bispira volutacornis, a dioecious polychaete, was followed over two spawning periods in Lettercallow Bay, on the west coast of Ireland. In the younger generation of sexually mature adults, gender can be determined on the basis of colour, the dorsal aspect of the males being cream at maturation, whilst the females are pink at the previtellogenic stage and an olive green at maturation. An annual reproductive cycle of 12 months shows that oocyte/sperm maturation is extraovarian. The male reproductive cycle proceeded at a more rapid pace than that of the female cycle, with sperm maturing earlier. Spawning in B. volutacornis occurred between late July and early September. B. volutacornis is a polytelic species, surviving spawning and releasing all gametes in a single batch, with a mean spawned oocyte diameter of 118 m. The reproductive cycle appears to be regulated by a seasonal change in water temperature, and spawning is timed to produce larvae during the autumnal planktonic bloom in Lettercallow Bay.Communicated by J.P. Thorpe, Port Erin     

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.     

Population dynamics and reproduction of Mysella bidentata (Bivalvia: Galeommatacea) in Galway Bay,Irish west coast

Mysella bidentata (Montagu) is found in high densities in an Amphiura filiformis (O. F. Müller) community in Inner Galway Bay, west coast of Ireland. As part of an ongoing investigation at one station within this community, the population dynamics and reproductive pattern of this bivalve were investigated during the period November 1978-December 1981. Monthly sampling over the 3 yr period revealed a mean population density of 745±672 individuals m^-2. Recruitment occurred during August-October and three year-classes were identified. Growth rates were assessed by analysis of length-frequency histograms and winter growth checks. Individuals from the Y1 and Y2 year-classes functioned as hermaphrodites during the 1980 and 1981 reproductive seasons. The Y0 year-class contained both males and hermaphrodites. Results are compared with available information on the same species from other locations.Contribution No. 225 from the School of Marine Sciences, University College, Galway Ireland     

Reproduction of<Emphasis Type="Italic"> Ophiactis resiliens</Emphasis> (Echinodermata: Ophiuroidea) in New South Wales with observations on recruitment

Reproduction of the ophiuroid Ophiactis resiliens Lyman 1879, a common species in the shallow waters around Australia, was examined over 1 year in two populations near Sydney, New South Wales. Gametogenesis was documented by histological examination of the gonads. Vitellogenesis in females and spermatocyte proliferation in males started in February and March. O. resiliens had an annual spawning period from May to September/October with peak gamete release from May to July. Thereafter, spawning occurred in some specimens through November. Gravid gonads were present for approximately 2 months longer in O. resiliens from Clovelly Bay than in those from Little Bay. At both sites spawning was episodic and gametogenesis continued through most of the spawning season. Most specimens (50–100%) collected from November to March had indeterminate/recovery stage gonads, the sex of which, could not be determined. The timing of gametogenic renewal during these months varied among individuals. O. resiliens has a 3-month planktonic stage and the presence of juveniles, 0.5 –1.5 mm disk diameter, in the coralline algal turf at Little Bay from February to May, suggests that settlement occurs in summer and autumn at this site. In contrast, juveniles were rarely encountered in coralline turf at Clovelly.     

Effect of inhibitors of carbonic anhydrase activity on photosynthesis in the red alga Soliera filiformis (Gigartinales: Rhodophyta)

The effect of light intensity, pH and carbonic anhydrase (CA) inhibitors on photosynthesis of the red marine macroalgae Solieria filiformis (Kützing) Gabrielson, collected from Taliarte (Gran Canaria, Canary Islands) in 1991, has been investigated. Plants taken from the sea (wild phenotype) developed spherical morphology (ball phenotype) after 2 mo culture in aerated tanks. The photosynthetic oxygen evolution in the wild phenotype was saturated at 100 mol photons m^-2s^-1, while the ball phenotype displayed saturation at 200 mol photons m^-2s^-1. The inhibitors of total CA activity (6-ethoxizolamide) and extracellular CA activity (dextran-bound sulfonamide) inhibited photosynthesis at pH 8.2, to 90 and 50% respectively, in both phenotypes. No inhibition of the photosynthetic oxygen evolution was detected at pH 6.5. CA activity was associated with both supernatant and pellet fractions of crude extracts of S. filiformis. The rate of alkalization of the medium by the algae was dependent on light intensity. We suggest that carbon dioxide is the general form of inorganic carbon transported across the plasmamembrane in S. filiformis. HCO₃ transport into the cell takes place simultaneously by an indirect mechanism (dehydration to CO₂ catalyzed by CAext) and by direct uptake. Extracellular (CAext) and intracellular (CAint) CAs are involved in the mechanisms of inorganic carbon assimilation by S. filiformis.     

Population dynamics and reproductive cycle of Pholoë minuta (Polychaeta: Sigalionidae) in Galway Bay

Pholoë minuta (Fabricius) is a prominent member of the Amphiura filiformis (O.F. Müller) subcommunity in Galway Bay, Ireland. The aim of the present investigation was to study the population dynamics and reproductive cycle of P. minuta in this area. Monthly sampling over a 2 yr period (November 1978–October 1980), at a permanent station (18 m) with a fine sand substrate, revealed a mean density of 96 individuals per 0.1 m². The width of the fifth elytron was correlated with total body length and was used as a basis for analysing population structure. Four distinct year classes were evident and estimates of recruitment levels and growth rates were calculated. P. minuta is polytelic and reproduces for the first time when approaching 3 yr of age. Spawning occurred in March—April and appeared to be associated with a seasonal low in bottom-water temperature.     

Gametogenesis in a Northumberland population of the polychaete Melinna cristata

The proliferative activity of the germinal epithelium in relation to breeding biology has been investigated in a population of the ampharetid polychaete Melinna cristata (Sars). This species is dioecious and highly gregarious, and breeds during a brief period at the end of December or beginning of January. The gonal exists in proliferative and non-proliferative states. The former begins immediately after spawning, and is maintained until September. The rate of release of gametocytes into the coelom varies during this period, with peaks in January and April. In September, DNA synthesis ceases in gonads of both males and females and no further gametocytes are released into the coelom, but primary oocytes which have completed the S phase of meiosis continue to develop and accumulate in the ovary as a population of previtellogenic oocytes which are released into the coelom immediately after spawning. This cycle of germinal epithelium activity is compared with those described in other polychaetes.     

The spatial structure of interzonal copepod populations in the Southern Ocean

The vertical distributions of Calanoides acutus, Calanus propinquus and Rhinicalanus gigas have been studied during Cruises I and II of the R.V. Ob in the Indian sector of the Southern Ocean. During the spawning period, all 3 species reveal the highest concentrations, and occupy the highest positions in the water column. During the period of feeding and growth, they begin slowly to disperse and to descend to greater depths. In spring, the populations ascend again to the water surface. In both seasons, later stages migrate first. Pronounced temperature gradients may prevent the ascent, and result in the breeding and subsequent growth of the new generation occurring in subsurface waters. Details of the vertical distributions of populations depend on their seasonal state and the local hydrology.     

Plasma sex steroid profiles and the seasonal reproductive cycle in male and female winter flounder,Pleuronectes americanus

The pattern of seasonal gonadal development and variations in plasma sex steroids were investigated in adult male and female winter flounder, Pleuronectes americanus (Walbaum), from Conception Bay Newfoundland beginning August 1987 to December 1988. The winter flounder reproductive cycle can be divided into five consecutive phases of relative reproductive activity including: (1) rapid gonadal recrudescence in the fall (August–December); (2) continued slow gondadal growth in females, or maintenance of the well developed gonads in males, during the winter (December–February); (3) a prespawning phase of gonadal maintenance in the spring (March–April); (4) spawning early in the summer (May–June) after the female gonads reach peak weight; and (5) the summer postspawned period (June–August) when the gonads remain regressed. Female gonadal recrudescence in August is characterized by small increases in plasma estrogen levels and recruitment of small oocytes (150 m) into yolk accumulation. For the winter months, estradiol-17 levels in the plasma remain stable, approximately 15 ng ml^-1, until rising again together with testosterone to peak hormone levels just prior to spawning in conjunction with the highest seasonal values for the gonadosomatic index (GSI) and oocyte diameter. After spawning, the ovaries are regressed and sex steroids in the plasma fall to very low levels. Rapid seasonal recrudescence of the gonads in males is evident from rising GSI values, which reach a maximum in October, and from substantial early seasonal increases in plasma testosterone and 11-ketotestosterone. Afterwards, although GSI values subsequently decline presumably reflecting the process of spermiogenesis and/or onset of spermiation in some males, the testes remain relatively well developed with the presence of sperm throughout the spawning season. As the proportion of spermiating males increases, the plasma levels of the androgenic steroid hormones rise to peak circulating levels from April to June. At the end of the summer reproductive season, the testes of post-spawned males become regressed and plasma testosterone and 11-ketotestosterone levels fall, reaching the lowest seasonal values.     

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.     

Tidal influences on the photoperiodic induction of tetrasporogenesis in Bonnemaisonia hamifera (Rhodophyta)

Photoperiodic control of tetrasporangium formation was studied in two low intertidal populations of Bonnemaisonia hamifera (Trailliella-phase) in Galway Bay, Ireland between September 1985 and February 1986. Photoperiodic induction was affected by tides, principally by the time of day at which high and low water occurred through the spring/neap cycle. High water of spring tides falling at the beginning and end of the day reduced the effective daylength for plants, and caused an early onset of reproduction. The threshold irradiance signalling light-dark transition was estimated to lie between (1-) 5 and 10 mol photons m^-2 s^-1 (400 to 700 nm). Low water of spring tides falling in the middle of the day exposed plants to high air temperatures during periods when seawater temperatures were below the lower limit for sporangium induction. This caused reproduction to be resumed. Quantitative differences between the two populations were caused by microclimatic effects (temperature, light reduction during tidal flooding) resulting from their different vertical heights on the shore.     

Population dynamics of Amphiura chiajei (Echinodermata: Ophiuroidea) in Killary Harbour,on the west coast of Ireland

The infaunal ophiuroid Amphiura chiajei Forbes is a dominant member of the A. chiajei community in Killary Harbour, a fjord-like inlet on the west coast of Ireland. High density populations (700 individuals/ m²) occur in sediments with a silt/clay content of 80 to 90% and organic carbon levels of 5 to 7%. A study of the population dynamics of this species was carried out from November 1985 to October 1988. At the outset, the population consisted of approximately equal numbers of adults and juveniles. By October 1988, the percentage of adults had increased to 95%, with little or no change in the mean annual dersities. However, there was some degree of variability between the mean monthly densities. Such variability may have been due to very low bottom temperatures, resulting in mortality among the older members of the population. Observations suggest that in Killary Harbour the species may attain an age of 10 yr, with an early annual growth rate of 0.5 mm (oral width). Annual recruitment was variable and low, due presumably to intraspecific competition with the adults; recruitment success may be very closely linked to adult mortality. The Killary Harbour findings are considered in the light of what is currently known of similar amphiurid populations in different geographical locations.     

Egg-capsule deposition and how behavioral interactions influence spawning rate in the squid<Emphasis Type="Italic"> Loligo opalescens</Emphasis> in Monterey Bay,California

Recent evidence suggests that, contrary to what was believed previously, most Loligo spp. females spawn multiple times and do not die immediately following a single spawning event. The present study used sustained focal observations of male/female pairs of the opalescent inshore squid Loligo opalescens Berry to examine the structure and behavior of near-bottom spawning groups. The study was carried out in a small area (10 km²) of Monterey Bay, California (36°36.1N; 121°53.4W), at depths of 25–45 m, using video cameras mounted on remotely operated vehicles. Behavioral observations were made primarily during daylight hours over known spawning beds in April and November 2000, and August 2001. Squid formed large aggregations in the water column where pairing occurred. Most commonly, only small numbers of active spawners were found at the substrate depositing egg capsules, and the mean operational sex ratio in the spawning groups was 1.87 males:1 female (range=1.0–8.5), although the ratio fluctuated rapidly as roving lone males joined and departed from the small spawning groups. On average, females (n=40) deposited 2.67 capsules (range 2–7) per focal observation at an average interval of 8.47 min between depositions (n=67). Following deposition of the capsules, females broke away from their consort males and jetted upwards to rejoin large schools located many meters above the substrate. Egg-capsule deposition was often interrupted by lone males seeking a mate, or by the approach of predators including fish and marine mammals. The results suggest that most of the communal egg beds in southern Monterey Bay are built up slowly through daily intermittent spawning, not in large big bang reproductive events as often depicted for L. opalescens.Communicated by J.P. Grassle, New Brunswick     

Photoperiodic activity pattern in the brittle star<Emphasis Type="Italic"> Amphiura filiformis</Emphasis>

Diurnal activity pattern of suspension-feeding arms of the infaunal brittle star Amphiura filiformis was recorded in situ by time-lapse photography at 30 m water depth in the Gullmarsfjord, west Sweden. Activity was clearly related to the photoperiod with high activity at night and low or no activity during daytime. In laboratory experiments, the activity of the arms of A. filiformis changed in relation to new manipulated photoperiods, demonstrating exogenous circadian rhythms. The time difference for the greatest change in activity of the arms between control and treatment was correlated with the photoperiodicity. The recently found microlenses and associated neural photoreceptors in ophiuroids may apply also to A. filiformis. The trade-off between efficient feeding and predator avoidance is discussed.Communicated by M. Kühl, Helsingør     

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     

Quantitative and ultrastructural studies on the reproductive biology of the polychaete Pholoe minuta in Galway Bay

Previous works by the present authors have identified, in a qualitative fashion, the reproductive cycle of Pholoe minuta (Fabricius) in inner Galway (53°13.5N; 9°7W). This quantitative, histological study, carried out over the period late May 1981 to late April 1982, shows that P. minuta is polytelic (two to three annual spawnings in an individual's lifespan). Individuals become sexually mature for the first time when approaching 3 yr of age. Estimates were made of the size of the reproductively active population (47 to 49%), sex ratio (1:1), duration (10 to 11 mo) and pattern (unimodal) of gametogenesis and fecundity levels (210 198 eggs m^-2). Maturation (mean oocyte size=85 m) and spawning (for 1 wk during late March through early April) were shown to be synchronised. Throughout oogenesis, follicle cells play an important role in binding together oocyte clusters, and desmosomal contacts are evident between oocyte and follicle cell plasmalemmas. No evidence for the synthesis of yolk precursors in the follicle cell cytoplasm was discovered. Oocytes are rich in Golgi apparatuses (yolk synthesising) and nuclear pores are numerous. Rough endoplasmic reticulum is distended in nature. Egg envelope development does not appear to commence until after the onset of vitellogenesis. Mature spermatozoa are of the primitive bullet-shaped type, with an unmodified acrosome. A possible method of elevation membrane formation is described.     

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.     

Annual reproductive cycle of the captive female Japanese sardineSardinops melanostictus: Relationship to ovarian development and serum levels of gonadal steroid hormones

Gonad and blood samples were taken from the captive female Japanese sardineSardinops melanostictus between 1988 and 1989, and changes in serum levels of gonadal steroids were correlated with the annual gonadal cycle. Under captive conditions, female fish did not mature and spawn spontaneously, although oocytes developed up to the end of vitellogenic growth. Based on evidence from ovarian histology, the annual gonadal cycle of the Japanese sardine was divisible into four periods, i.e., immature (June to October), vitellogenesis (November to December), spawning (January to March), and post-spawning (April to May). The pattern of seasonal change in the gonadosomatic index (GSI) showed an inverse correlation to change in water temperature and reflected the degree of ovarian maturity. The serum estradiol-17 level increased from its lowest concentration (0.12 ng ml^–1) in September to a peak (1.14 ng ml^–1) in March. Serum 17,20-dihydroxy-4-pregnen-3-one (17,20-P) was detectable at low levels (<0.3 ng ml^–1) between October and February, but was below the assay detection limit (0.06 ng ml^–1) at all other times. Testosterone was not detectable (<0.06 ng ml^–1) in the serum of any fish throughout the year. The effects of several steroids on the maturation of follicle-enclosed oocytes of sardine were examined in vitro, and 17,20-P was found to be the most potent inducer of maturation. This suggests that post-vitellogenic oocytes of the Japanese sardine in captivity have an ability to respond to an appropriate hormonal effector and subsequently to resume meiotic maturation.