Seasonal changes in the biochemical composition of the chiton Chiton iatricus (Polyplacophora: Mollusca) and the marine pulmonate Onchidium verruculatum (Gastropoda: Mollusca) in relation to their reproductive cycles

Seasonal changes in the biochemical composition of two molluses, gonochoric Chiton iatricus and a marine pulmonate, Ochnidium verruculatum inhabiting the same tropical area, but having different modes of reproduction, were studied. Protein is the major biochemical component stored in C. iatricus, whereas lipid is predominantly stored in O. verruculatum. The correlation between these quantifications and the annual reproductive cycles of the two species investigated suggest that nutrients stored in the somatic body parts are channeled towards the gonad during the gametogenesis. In C. iatricus sexual differences efrist in the biochemical needs of the ovary and the testis. The ovary always has a higher protein content than the testis. In the hermaphrodite pulmonate O. verruculatum, biochemical expenditure on gamete production is relatively low because this species has to spend energy on the reproductive propagation viz. copulation and egg-laying. On the contrary, in C. iatricus the biochemical energy is mainly utilized for the production of gametes which have to be produced in great numbers in order to counter-balance the loss of gametes caused by external fertilization. The collected data are discussed in the light of comparative physiology of reproduction.     

Annual reproductive cycle of the chiton Acanthopleura japonica

Annual reproductive cycle of the chiton Acanthopleura japonica (Lischke) was studied by observing both gonad index and histological features from October 1980 to October 1981. No significant changes in each factor were observed from October to April. Increased values of gonad index and spermatogenesis or vitellogenesis in gondas were observed from May to September. In the same time, gonad index decreased and sperms or oocytes in gonalds disappeared in the days around spring tide, and they were recovered around neap tide. Suggested spawning around spring tide from May to September is also supported by the occurrence of released eggs in the days around the new and full moons.     

Reproductive synchrony in a habitat-forming kelp and its relationship with environmental conditions

Key biological processes such as the timing and synchrony of reproduction, are sensitive to fluctuations in the environment, as well as alterations in the physiology and behaviour of individuals and the ecology of populations. The main aim of this study was to identify patterns in reproductive timing and relationships with prevailing environmental conditions, for the major habitat-forming alga in temperate Australia, Ecklonia radiata (C. Agardh) J. Agardh. We observed strong synchronous patterns of zoospore production across spatial scales from hundreds of metres to tens of kilometres, with populations of E. radiata reproducing from mid-summer to the end of autumn. High zoospore densities were positively related to sea temperature and E. radiata released zoospores in temperatures well above previously documented thermal tolerance limits for other Laminariales. Reproductive timing was also negatively related to seasonal increase in wave exposure. We conclude that E. radiata undergoes synchronous reproduction which can be linked to its annual growth cycle and natural environmental fluctuations, to promote growth and survival of recruits.     

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.     

Energy expended on growth and gonad output in the ribbed musselAulacomya ater

Length/weight relationships have been computed for shell, flesh and byssus ofAulacomya ater (Molina) and energy values used to convert the weights to energy equivalents. Shell accounts for some 26% of total body energy, while the contribution of the byssus declines from 15 to 8% during growth. Observations of juvenile growth rates have been used to generate a Gompertz growth equation which predicts attainment of maximum length (90 mm) after 11 years. Reproductive condition has been assessed by monitoring seasonal fluctuations in the flesh weight of standard-sized individuals, calculated from monthly length/weight regressions. There appear to be three spawnings, of variable date and intensity, each year. From the above data, annual energy expenditure on growth and gonad output has been calculated for individuals of various sizes. The ratio of total production to biomass is a declining clining function of shell length, dropping from 29.5 at 5 mm to 0.8 at 85 mm. The proportion of total production expended on gamete output increases steadily from 25% at attainment of maturity (15 mm) to 81% at 85 mm length. The considerable effects of changing size composition on the amount and type of production in natural populations are discussed.     

Reproductive cycle and seasonal feeding activity of the neogastropod Buccinum undatum

The whelk Buccinum undatum L. in the northern Gulf of St. Lawrence has a distinct annual reproductive cycle. The ovary and pallial oviduct of the female, and the seminal vesicle of the male, show parallel development, whereas the testis has an inverse cycle. Thus, at the end of winter when the ovary, pallial oviduct and seminal vesicle reach maximal size, the testis is small and undeveloped, whereas the reverse situation occurs in summer. This is the first report of a totally inverse relationship between the ovary and testis for a marine invertebrate. This unusual pattern is due to the long-term storage of sperm in the seminal vesicle of the male prior to copulation. There is no resting period at the end of oogenesis, but rather renewed ovarian development starts shortly after gamete release. By contrast, in the testis after transfer of sperm to the seminal vesicle at the end of spermatogenesis, there is a distinct period of phagocytosis. Mating begins in mid-May, reaches a peak during June and terminates prior to July. Egg laying begins in late May, reaches a peak in mid-June to mid-July and can continue until late August. Examination of the incidence of individuals with food in their stomachs showed that feeding activity is maximal in late autumn through to early spring and decreases sharply coincident with the onset of breeding in late May.     

Reproductive cycle of the brachiopodTerebratulina retusa on the west coast of Scotland

The reproductive cycles of two populations of the Recent articulate brachiopodTerebratulina retusa (Linnaeus) are described, based on samples taken between October 1985 and October 1986. Stereological analysis of the gonads revealed a single synchronised spawning event between late November and the end of January in the Firth of Lorn. In a second population, in Loch Fyne, spawning took place repeatedly throughout the spring and summer, with greatest spawning activity occurring in the late autumn.T. retusa from Loch Fyne were approximately three times more fecund than those from the Firth of Lorn. Productivity in the Clyde Sea area is approximately five times higher than in Firth of Lorn and higher fecundities and spawning frequencies of the Loch Fyne population are attributed, in part, to greater food availability. The initiation of gametogenesis appears to be mediated by the mobilisation of reserves stored in the outer mantle epithelium during the winter. Primary productivity increases dramatically in the spring and gametogenesis shows a corresponding increase.     

Seasonal changes in pyloric caecum and gonad indices during the annual reproductive cycle in the seastar Asterias forbesi

Seasonal changes in pyloric caecum and gonad indices were studied in a population of the seastar Asterias forbesi (Desor) from East Rockaway Inlet (Long Island, New York) during two annual cycles (1980/81, 1982/83). Pyloric caeca indices increased during fall, reached a maximum about April, and declined sharply to a minimum in mid-summer. Gonad indices increased during fall and winter and reached a maximum about May. Judging from gonad size analysis, spawning occurred in late June to early July, at bottom temperatures of 16° to 18°C. There was no long-term inverse relationship between pyloric caecum and gonad indices. Since nutrients and energy ingested during the important fall feeding period are utilized simultaneously for body growth and gonad development, it is unnecessary for the pyloric caeca to store nutrients over long periods as occurs in A. rubens and many other species. The specific caeca-gonad relationship is probably an adaptation to the extreme seasonal thermal fluctuations of the NW Atlantic, which constrain the feeding activity of A. forbesi and limit the possibility of long-term storage. In this population, most individuals grow rapidly in their first year (including first two summers) and spawn in their third summer. Few seastars survive to spawn again in their fourth summer. However, gametogenesis may take place in some individuals in their second summer. Year-to-year variability in mean size at spawning may reflect temporal variation in environmental conditions (weather, food availability) at this site.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Trail following in the chiton Acanthopleura gemmata: operational and ecological problems

The Indo-Pacific, intertidal chiton Acanthopleura gemmata (Blainville) is a central place forager which occupies definite, actively dug and defended scars, and migrates during nocturnal low tides up to 3 m away to feed on algal grounds. After each feeding excursion, most chitons home precisely to their scars. Photographic tracking, using lightemitting diodes (LED) glued to the chitons, and experiments were conducted throughout 45 d (August to September 1984) along the Nimu Peninsula (Somalia) in order to elucidate the importance of trail following in the homing and feeding of this species. LED tracking of the chitons throughout each activity phase and trail interruption experiments showed that following the personal outgoing trail is a basic homing mechanism in this species. Translocation of the chitons on a conspecific trail and homing performance analysis revealed that the trail-associated information involved in the homing is polymorphic in the population, thus minimizing the probability of following a conspecific trail despite the frequent trail crossings due to high population density. The LED tracking throughout successive activity phases showed a high coincidence between the paths traced by the same chiton night after night. In the few cases when the outgoing and return paths of the first night markedly diverge, the home-feeding ground route of the successive night overlaps the return trail of the previous night. A less pronounced inter-individual trail following also occurs during foodward excursions. These findings suggest that the trails released by A. gemmata are complex, including both quasi-individual and species-specific information. Moreover, this chiton may utilize its trail following capacity not only to home, but also for the retrieval of the feeding grounds night after night.     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

The larval settling response of the lined chiton Tonicella lineata

The lined chiton Tonicella lineata (Wood, 1815) is found on enerusting coralline algae in the lower rocky intertidal zone of Oregon and San Juan Island, Washington, USA. An encrusting coralline alga is the major food item of this chiton. Experiments were performed to test the settling response of T. lineata larvae to various algae and other substrata. In these experiments, the larvae would settle only on pieces of encrusting coralline algae and piecesof ceramic roofing tile soaked in a coralline algal extract. The settling stimulus is probably chemical in nature, and is inactivated by boiling. In laboratory cultures, normal development stops at the trochophore stage (110 to 160 h, depending on the temperature). Metamorphosis and further development will take place only after stttlement on encrusting coralline algae.     

Quantitative studies on gonad cycle,fecundity, reproductive output and recruitment in a baltic Mytilus edulis population

Since Mytilus edulis L. is a biomass dominant in the Baltic much interest is focused on the ecology of the species. In this paper an attempt is made to quantitatively cover the reproductive cycle of a Baltic M. edulis population in order to provide data for energy flow models and to discuss aspects of recruitment in this species. Histological preparations of gonads showed that gametogenesis started with declining temperatures in autumn and proceeded very slowly through winter. At the beginning of March when food was supplied during the spring phytoplankton bloom, rapid maturation took place. This was also revealed by an increase in meat weight of the mussels. Only one spawning period was recorded, from the middle of May until the beginning of June, due to food being strongly limited to the population during the rest of the year. The length of the larval period was estimated as being 5 to 6 wk and settlement was registered from the end of June through July. In general the large annual variations found in the Baltic with regard to temperature and food abundance give rise to a more marked annual pattern in the reproductive cycle than is encountered in other seas. Fecundity was assessed for two populations from 4-and 15-m depths from studies covering two annual cycles of the changes in the relation of shell length — meat weight. The size-related fecundity was found to be equal in both populations and related to food abundance and not to growth or age. Fecundity, expressed as weight loss at spawning, ranged from 0% in 2-mm mussels and increased from 38 to 52% in 10-to 30-mm mussels. The fecundity as percentage of biomass in full-grown Baltic M. edulis is of similar magnitude as in full-grown mussels from other areas despite the smaller size of Baltic mussels. The reproductive output for the total 160 km² research area was calculated as being 1 200 tons dry weight or 80% of the standing stock, which, due to the particular features of the Baltic M. edulis population probably represents the larger part of the total mussel production. This reproductive output, calculated as 8·10⁷ eggs·m^-2 and corresponding to 50% of the total annual zooplankton production, may thus consitute an important food source for herring larvae and carnivorous zooplankton. Recruitment was divided into two phases: (1) Recruitment of juveniles (=settlement of larvae), and (2) recruitment to breeding stock. Monitoring studies of settlement on ropes and the year round presence of high abundances of mussels <2 mm indicate that settlement is in excess of the demands for maintaining population size and that most settled mussels form a pool of competitively suppressed non-growing individuals. Not until death of an already established mussel will these become recruited to the breeding population. Thus recruitment is possible throughout the year which stabilizes the population and maintains it near the carrying capacity of the area with regard to food and space availability.     

Economic growth and environmental pollution under the max-min principle

This paper first analyzes the interactions between capital accumulation and environmental pollution under the program of sustained constant utility, which depends on per capita consumption and pollution, over all generations. Then the optimal program of economic growth is discussed. The implications of the constant-utility criterion (or the max-min principle) are compared with those of the utilitarian criterion which has been widely adopted in environmental pollution literature. Subsequently, the analysis incorporates pollution-abatement investment and exhaustible resources. We conclude that the max-min principle works fairly satisfactorily in this world.     

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.     

Effects of temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis

To assess the effects of both temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), individuals collected December 1996 (winter experiment) and June 1997 (summer experiment) were maintained for 3 months in one of four experimental treatments: (1) 3 °C and fed ad libitum (high ration), (2) 3 °C and fed one-seventh of the maximum ration (low ration), and (3) 12 °C and fed the high ration; (4) 12 °C and fed the low ration. All individuals were fed an artificial diet and exposed to only 1 h of light every day. At the end of both experiments, mean gonad indices of sea urchins fed the high ration had increased significantly (11–24% and 6–19% in the winter and summer experiments, respectively), while the gonad indices of individuals fed the low ration did not change. At the high ration (both experiments), the increase in gonad index of sea urchins occurred primarily as the result of a significant increase in the mass of nutritive phagocytes, as revealed by histological analyses. Primary oocytes were significantly larger in individuals held at 3 °C than at 12 °C throughout the winter experiment, regardless of food ration; during the summer experiment, primary oocytes were significantly larger in individuals receiving the high ration, regardless of the temperature at which they were held. These results suggest that: (1) food availability is the most important factor regulating energy storage and the relative size of gonads throughout the year, (2) temperature affects the rate of growth and maturation of primary oocytes during the later stages of oogenesis, and (3) once gametogenesis has been initiated, mature ova will be produced, even under conditions of low food availability. Conditions of high food availability in summer and low temperature in winter would thus favor reproductive output in sea urchin populations. Received: 1 March 2000 / Accepted: 4 October 2000     

Reproductive cycle,development and recruitment of two geographically separated populations of the dendrochirote holothurian Aslia lefevrei

The boreo-mediterranean holothurian Aslia lefevrei (Barrois) occupies cryptic habitats from low-water mark to ca. 20 m. During the present work specimens were collected from the west coast of Ireland (February 1979 to August 1980), where A. lefevreiis a prominent element in two discrete epibenthic assemblages, and from the north-west of Spain (September 1981 to August 1982), where this holothurian is densely aggregated in association with raft cultivation of the mussel Mytilus edulis. Its reproductive cycle, spawning process, embryological development, settlement, and early growth rates have been documented by means of laboratory and field studies. Depending on geographic location, spawning in the wild occurs between January and May. Development to the gastrula takes some 3 to 4 d. Uniformly ciliated dipleurula/auricularia hatch 12 to 24 h after gastrulation. The pentactula stage is attained approximately 7 d after fertilization, following which the larva settles. Newly settled individuals utilize the five primary tentacles for both feeding and locomotion. With the advent of small numbers of podia in the ambulacral regions, two of the primary tentacles prolong their locomotory and anchorage capabilities while the remaining tentacles develop adult characteristics and function solely in feeding. Growth was followed for an estimated 30 mo period using length-frequency analyses. In the first year of life, A. lefevrei attain an average length of 3.4 cm; individuals on the sea floor can attain up to 18 cm, indicating a possible life expectancy of 10 yr or more.Contribution No. 301 from the School of Marine Sciences, University College Galway, Ireland     

Reproductive cycle of Zidona dufresnei (Caenogastropoda: Volutidae) from the southwestern Atlantic Ocean

Gonadal development of Zidona dufresnei (Donovan, 1823) (Caenogastropoda: Volutidae) was studied over a period of two consecutive years, through analysis of gonadal tissue. Individuals were sampled monthly at the Mar del Plata area, Argentina. Gonadosomatic index was estimated for males and oocyte size was used to estimate the stage of gonadal development in females. The reproductive season in this species in the sampled locality extended from October to March (austral spring-summer). During summer, a stage of advanced gonadal development and spawning predominated in the adult population. In autumn, gonads were generally under atresia; during wintertime (June-August), they underwent a period of recovery that lasted until the spring months, when gametes were released again. Synchronism between both sexes was evident. Marked periods of spawning were followed by resorption periods and then a growing phase; it was very clear that reproductive seasonality was linked to changes in bottom water temperature. These results suggest that Z. dufresnei gonads have a yearly cycle of gamete production, with two major activity peaks in September-October and January-February.     

Genetic and nutritional control of the gametogenic cycle in Mytilus edulis

Individuals of Mytilus edulis were collected from two populations in easterm Long Island Sound, USA, at 6 wk intervals from January 1982 to February 1983, and the hypothesis that nutrient availability controls the timing of gametogenic events in this mussel was tested using genetic variants at the Lap locus. In eastern Long Island Sound, Lap genotypes differ in the net rate of nitrogen accumulation. A highly significant difference in reproductive cycle among genotypes occurred in a population of M. edulis where Lap genotypes differ in nitrogen budget. Genotypes with lower rates of nutrient accumulation delayed the reproductive cycle by approximately six weeks relative to genotypes with high rates of nutrient accumulation. No significant difference in reproductive cycle occurred at a locality where Lap genotypes do not differ in nitrogen budget. These data indicate that nutritional status can exert significant control over the timing of gametogenesis in M. edulis. Differences in reproductive cycle among Lap genotypes provide a mechanism of non-random mating among genotypes which may result in heterozygote deficiencies at this locus.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.