Seasonal cycle of gonadal development and serum levels of vitellogenin of the red porgy, Pagrus pagrus (Teleostei: Sparidae)

Histological examination of gonads of female and male red porgy, Pagrus pagrus, reared in captivity, was carried out in order to describe the main gonadal changes related to gametogenesis and the seasonal changes of environmental factors. The gonadosomatic index (GSI) and the concentration of serum vitellogenin (VTG) were also determined. The frequency distribution of gonad development stages and the GSI and vitellogenin concentration during the annual cycle indicated the separation of the female and male reproductive cycles into three main periods. The autumn period when gametogenesis begins (October-November), the period of exogenous vitellogenesis (December-March) or spermiation (December-March), and the spawning season (March-May). The spawning period coincided with an increase in temperature (15-19°C) and daylength. Serum levels of vitellogenin rose significantly in January, reached a peak at the beginning of the spawning period (March, 405.5 µg ml^-1) and remained high until the end of the spawning period (May).     

Changes in the content and fatty acid composition of lipid in an isolated population of the capelin Mallotus villosus during sexual maturation and spawning

Changes in the wet weights and lipid contents of muscle, liver and gonad were determined in male and female Mallotus villosus in Balsfjorden, northern Norway, in 1981, from January, when gonadal development becomes noticeable, until May when the fish are spawning. Fatty acid compositions of tissue lipids were also determined. Over 4 mo prior to spawning, the weight of muscle in female capelin decreased by 32% while the weight of the ovary increased exponentially by 830%. In males the weight of the muscle remained constant and that of the testis decreased slightly. The lipid contents of the muscle of both males and females decreased by 76% over the period and an inverse relationship existed between the water and lipid contents of muscle in both sexes. Male liver weight remained constant over the period of study whereas female liver weight increased transiently by 300% between January and March. 38% of the lipid lost from female muscle was accounted for by lipid deposited in ovary whereas negligible amounts of the lipid lost from male muscle was accounted for by lipid in the testis. Gonadal lipid was always richer in polyunsaturated fatty acids than muscle lipid and, immediately prior to spawning, 42% of the fatty acids in ovarian lipid were polyunsaturated. Muscle lipid of males and females showed a progressive increase in the percentage of the long-chain monoenes 20:1 and 22:1 between January and May. It is concluded that male capelin catabolise more of their muscle lipid reserves than females in the 4 mo prior to spawning and that most of the lipid catabolism in males is associated with physical activity. Conversely, females deposit much more of their muscle lipid in gonads than males, although considerable selectivity occurs in the mobilisation of fatty acids from muscle lipid into ovarian lipid. Additionally, biosynthesis of gonadal constituents accounts for a considerable proportion of the lipid catabolised in females.     

Egg production and lipid content of Calanus glacialis in spring: indication of a food-dependent and food-independent reproductive mode

Female Calanus glacialis were collected in early May 1989 in the pack ice region of the western Barents Sea and were fed or starved over 11 wk. Both groups laid eggs continuously during this period, however, fed females laid up to six times more eggs. During the first 10 d after collection, both groups spawned at low rates. There-after, fed females strongly increased spawning rates and maintained high egg production levels over 11 wk, while the rates of starved females decreased. During starvation they lost 70% body carbon, 50% body nitrogen and 70% lipids. The wax ester portion decreased from 86 to ca. 60% of total lipids. Three phases of gonad development and lipid metabolism were distinguished: early gonad development; gonad maturation with a rapid decrease in lipids, especially wax esters; and spawning under fed and starved conditions, where in fed females food provided most of the energy, whereas in starved females the lipid content strongly decreased.     

Effects of seasonality and population density on the reproduction of the Indo-Pacific echinoid<Emphasis Type="Italic"> Echinometra mathaei</Emphasis> in Kenyan coral reef lagoons

Reproduction in the widely distributed tropical sea urchin Echinometra mathaei (de Blainville) was examined in three Kenyan reef lagoons that differed in substrate cover and E. mathaei population density. Histological examination of the gonads and gonad index measurements for 3 years showed a seasonal reproductive cycle with gametogenesis commencing in July, when temperature and light are at their lowest, and spawning commencing in December and peaking in February to May, when temperature and light reach their annual maxima. Monthly gonad indices correlated significantly with seawater temperature and light intensity. Male urchins had mature gametes for a longer period (8 months) than females (6 months), possibly an adaptive strategy that increases the probability of fertilization. Lunar periodicity was not observed, as male and female gonads were full of gametes on all days of the lunar cycle during the period of spawning. The peak in spawning activity coincides with the peak in phytoplankton abundance, which could ensure high food availability for the planktonic larvae. Gonad weights were significantly higher relative to urchin weight at the reef having the largest mean sizes and lowest population of urchins, indicating availability of food resources for growth and reproduction. E. mathaei at the reef with the highest density had the smallest urchins and high relative gonad sizes, indicative of a tradeoff between growth and reproduction when food is limited.Communicated by O. Kinne, Oldendorf/Luhe     

The gametogenic cycle of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Mollusca: Cephalopoda) as observed on the Atlantic coast of Andalusia (south of Spain)

The present study reports the gametogenic cycle of the common octopus, Octopus vulgaris Cuvier, 1797 in the Atlantic coast of Andalusia between May 2001 and May 2002. A total of 673 specimens of total weight ranging from 30 to 3,732 g were analysed. The gonad development was defined by means of histological methods and gonadosomatic indexes. Four gametogenic stages were identified in males (I immature, II maturing, III mature, IV spawning) and in females (I immature, II maturing, III pre-spawning, IV spawning). The sex ratio obtained was not significantly different from 1:1 (P>0.01). Mature males were found during the whole year, September being the month when the highest rate of sexually active individuals occurred (48%). In females, the gametogenic cycle commenced in December and ended in November, and spawning occurred between April and October, with two maximum peaks in August and September (48% in each). Size at sexual maturity was 850 g of total body weight for males and 1,250 g of total body weight for females. Males mature earlier than females.Communicated by S.A. Poulet, Roscoff     

Energy storage and dynamics in bay anchovy Anchoa mitchilli

The energy of the body components and the energy costs of spawning and overwintering in the bay anchovy Anchoa mitchilli, the most abundant fish in Chesapeake Bay, were studied to determine seasonal variability during the different stages of its life cycle. Bimonthly samples were collected by trawl from April 1990 through October 1991. Fish condition and body energy levels fluctuated seasonally, and were related to anchovy size. Energy equivalents (cal g^-1 dry wt) was highest in December, before the overwintering period. The somatic weight component increased by 32 to 33% and total body weight by 26% during the spawning season, indicating that feeding not only met energy requirements of daily spawning but also provided surplus energy for growth. The overwintering loss of energy was 33 to 35% of total body calories, and was primarily derived from deposit fat in somatic and visceral tissues.     

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.     

Seasonality of feeding and nutritional status during the austral winter in the Antarctic sea urchin Sterechinus neumayeri

The seasonal pattern of food intake and tissue energy status was measured over a 2 year period for two populations of the common Antarctic echinoid Sterechinus neumayeri living on contrasting substrata at Rothera Point, Adelaide Island, Antarctica. Food intake was estimated from faecal egestion, and both tissue mass and energy content assessed. Food availability was intensely seasonal, in that water column chlorophyll content and sediment pigment content varied markedly throughout the year. In response, both urchin populations showed an extremely seasonal cycle of faecal egestion, indicating a strong seasonality of feeding activity. Urchins from North Cove living on soft sediment fed at a considerably higher rate, and had a significantly larger Aristotle's lantern, than those in South Cove, 1 km away, living on hard substrata and taking a more cosmopolitan diet. Faecal egestion in both populations was zero for a 7 month period in the austral winter of 1997, and again for a 4 month period in 1998. During the austral summer of 1997/1998, 84% of the total annual energy intake took place in the period January to March. Significant decrease in gut tissue mass provided energy for maintenance in early winter, although progressive reduction of both gonad energy content (but without a detectable change in gonad mass) and body wall organic mass provided energy during the late winter period. The mass of reproductive tissue showed large differences between the two sites, but there was no marked decrease associated with spawning. Tissue proximate composition was assessed stoichiometrically from elemental composition and also checked by direct assay; gonad tissue was richer in lipid than gut tissue, though both were dominated by protein and contained only small amounts of carbohydrate. These data suggest that the very strong seasonality of food intake does not pose a significant energetic challenge for this species.     

Coexistence and reproductive isolation of the sympatric echinoids<Emphasis Type="Italic"> Diadema savignyi</Emphasis> Michelin and<Emphasis Type="Italic"> Diadema setosum</Emphasis> (Leske) on Kenyan coral reefs

The sympatric echinoids Diadema savignyi and D. setosum coexist in shallow reef lagoons throughout East Africa. The reproductive strategies of these echinoids were studied to investigate reproductive isolation as a possible mechanism for maintaining the coexistence of these closely related species. The annual reproductive cycle and lunar periodicity were determined by gonad index measurements, histological examination of gametogenesis, and induction of spawning with injections of KCl. The peak reproductive period of D. savignyi coincided with the north-east monsoon period (when light and temperatures are high) as gonad indices were high (>8%) beginning in February and peaked at 9.7% in May. Gonad indices subsequently rapidly decreased (by 26%) in June at the beginning of the cooler south-east monsoon period. However, the presence of sperm and ova in most months of the year indicates continuous gametogenesis with reduced reproductive effort during the cooler months. The annual cycle for D. setosum showed less of a seasonal trend as gonad indices remained above 7% throughout much of the year but tended to be highest when temperatures were lower. This is the first confirmation of continuous reproduction in these two species at the equator. The reproductive patterns of both species remained consistent over 2 years of sampling. Both species exhibited a synchronized lunar spawning periodicity during the 3 months sampled, with D. setosum spawning on lunar days 8–10 and D. savignyi spawning after the full moon (lunar days 17–18). Whereas spawning in D. savignyi was very tightly synchronized, 20% of D. setosum individuals still spawned after the peak spawning period. The coexistence of these closely related species appears to be maintained by temporal reproductive isolation during the lunar spawning period reinforced by seasonal differences in reproductive effort.Communicated by O. Kinne, Oldendorf/Luhe     

Effect of sexual maturation on the tissue biochemical composition of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> and<Emphasis Type="Italic"> O</Emphasis>.<Emphasis Type="Italic"> defilippi</Emphasis> (Mollusca: Cephalopoda)

Changes in the protein, lipid, glycogen, cholesterol and energy contents, total amino acid and fatty acid profiles of Octopus vulgaris and O. defilippi tissues (gonad, digestive gland and muscle) during sexual maturation (spermatogenesis and oogenesis) were investigated. Both species showed an increase of amino acids and protein content in the gonad throughout sexual maturation (namely in oogenesis), but allocation of these nitrogen compounds from the digestive gland and muscle was not evident. The major essential amino acids in the three tissues were leucine, lysine and arginine. The major non-essential amino acids were glutamic acid, aspartic acid and alanine. With respect to carbon compounds, a significant increasing trend (P<0.05) in the lipid and fatty acid contents in the three tissues was observed, and, consequently, there was also little evidence of accumulated lipid storage reserves being used for egg production. It seems that for egg production both Octopus species use energy directly from food, rather than from stored products. This direct acquisition model contrasts with the previous model for Octopus vulgaris proposed by ODor and Wells (1978: J Exp Biol 77:15–31). Most of saturated fatty acid content of the three tissues was presented as 16:0 and 18:0, monounsaturated fatty acid content as 18:1 and 20:1 and polyunsaturated fatty acid content as arachidonic acid (20:4n-6), eicosapentaenoic acid (20:5n-3) and docosahexaenoic acid (22:6n-3). Though cholesterol is an important precursor of steroid hormones, this sterol content exhibited variations that do not seem to be related with the maturation process. Moreover, significant differences (P<0.05) were obtained between genders, suggesting that perhaps there is a greater physiological demand for cholesterol during spermatogenesis than oogenesis. If the component sterols of octopus are of a dietary origin, considerable variation in the cholesterol content between species might be expected on the basis of the sterol composition of their prey. The glycogen reserves increased significantly in the gonad and decreased significantly (P<0.05) in the digestive gland and muscle of O. vulgaris (these trends were not evident in O. defilippi). Glycogen may play an important role in the maturation process and embryogenesis of these organisms, because carbohydrates are precursors of metabolic intermediates in the production of energy. It was evident that sexual maturation had a significant effect upon the gonad energy content, but the non-significant energy variation (P>0.05) in the digestive gland and muscle revealed no evidence that storage reserves are transferred from tissue to tissue. The biochemical composition of digestive gland and muscle may not be influenced by sexual maturation, but rather by other biotic factors, such as feeding activity, food availability, spawning and brooding.Communicated by S.A. Poulet, Roscoff     

Gonad morphology and oocyte development in<Emphasis Type="Italic"> Pseudocalanus</Emphasis> spp. in relation to spawning activity

Gonad maturation processes were studied in Pseudocalanus spp. females from Georges Bank and Cape Cod Bay (northwest Atlantic) using a combination of morphological analysis and experiments. For light microscopy of the oocytes, females of different maturation stages were preserved immediately after capture. The maturation processes during the spawning cycle were described from observations of live females which were exposed to feeding and starvation at two temperatures, 8 and 15°C, for 12 days. The gonad morphology of these females was examined in 24 h intervals, and spawning events were recorded. Both light microscopy and whole animal observation revealed that oocytes during maturation change in shape and size, in the morphology of the nucleus, and in the appearance of the ooplasm. Due to these modifications of oocyte morphology and due to oocyte migration, the morphology of the gonads changed distinctly during a spawning cycle. Five oocyte development stages were identified by light microscopy and related to a macroscopic system of four gonad development stages, that can be applied to whole animals and allows the identification of females ready to spawn. The experiment showed that food and temperature had strong effects on gonad maturation processes. High proportions of mature females were found when food was available, whereas the proportion of immature females increased shortly after exposure to starvation. Compared to 15°C, gonad maturation at 8°C was prolonged and thus spawning frequency was lower. The final maturation processes at food saturated conditions were slower than the embryonic development, and no indication was found that mature oocytes are stored in the diverticula waiting to be released. The duration of the interspawning interval would thus be determined by the duration of final oocyte maturation which is dependent on both temperature and food supply.Communicated by O. Kinne, Oldendorf/Luhe     

Changes in dopamine,norepinephrine and serotonin levels in the pituitary,telencephalon and hypothalamus during gonadal development of male <Emphasis Type="Italic">Lutjanus argentiventris</Emphasis> (Teleostei)

Biogenic amines are important in the regulation of certain aspects of teleost reproduction, but relatively little is known about variations in their levels in the nervous system during the reproductive cycle. The aim of this study was to investigate changes in the levels of dopamine (DA), norepinephrine (NE), serotonin (5-HT), the DA metabolite 3,4-dihydroxyphenylacetic acid (DOPAC) and the 5-HT metabolite 5-hydroxyindoleacetic acid (5-HIAA) in the pituitary and brain (hypothalamus and telencephalon) of the adult male of the tropical marine yellow snapper Lutjanus argentiventris during the reproductive cycle (quiescent, prespawning and spawning periods). Gonad weight, gonad/body weight ratio and plasma-testosterone levels increased steadily throughout gonadal development, but 17#-estradiol levels did not vary significantly. Pituitary DA and DOPAC levels and the DOPAC/DA ratio declined with gonadal development, whereas hypothalamic DOPAC level and DOPAC/DA ratio increased. In the telencephalon, DA and DOPAC levels and DOPAC/DA ratio all peaked in the prespawning period; however, DA level was lowest in the spawning period, whereas DOPAC level and DOPAC/DA ratio were both lowest in the quiescent period. Telencephalic NE, 5-HT and 5-HIAA levels likewise all peaked in the prespawning period, and were lowest in the spawning period. Telencephalic 5-HIAA/5-HT ratio peaked in the quiescent period and was lowest in the spawning period. No significant variation was observed either in pituitary NE level or in hypothalamic NE, 5-HT and 5-HIAA levels. The present results indicate that males of L. argentiventris show a decline in pituitary DA activity during gonadal development. We discuss the possible involvement of biogenic amines in the regulation of GTH II secretion, through direct action on the pituitary or indirect modulation of gonadotrophin-releasing hormone neuron activity.     

Effect of diet on growth rate and reproductive fitness of Turbo sarmaticus (Mollusca: Vetigastropoda: Turbinidae)

Experiments were conducted to determine the effect of four algal diets (Corallina spp., Gelidium pristoides, Ulva rigida and an equal volume mixture of these seaweeds) on growth and reproduction of the South African gastropod Turbo sarmaticus Linnaeus, 1758. The best growth rate of juveniles (up to 13.8 mm shell length and 34.26 g wet body weight increase in 12 months), reproductive fitness of mature specimens (gonad index up to 33%) and energy reserves (up to 4.76 mg glycogen/100 mg foot tissue) were achieved when T. sarmaticus was fed U. rigida or a mixed diet. In addition, the gonad index of individuals fed these diets was almost twice that of similar-sized field specimens. Juveniles fed Corallina spp. only, grew very little (only 2.4 mm shell length and 4.23 g wet body weight increase in 12 months). The reproductive fitness of adults fed on such a diet was also poor (gonad index <4.5%) and energy reserves were low (<3.5 mg glycogen/100 mg foot tissue). Along the southeastern coast of South Africa, T. sarmaticus has a distinct reproductive cycle with gametogenesis occurring from March/April until August/September, whilst maturity (gonad index = 15%) was maintained until December, after which spawning occurred until March. Received: 6 July 1998 / Accepted: 8 March 1999     

Sub-lethal impact of carbaryl on food utilization in the freshwater prawn Macrobrachium malcolmsonii

This study determines the toxic effect of carbaryl (Sevin50% W.P) on the food utilization parameters in intermoult juveniles of the prawn, Macrobrachium malcolmsonii. The prawns (4.5-5.0 cm in length and 1.0-1.25 g wet wt.) were exposed to three sub-lethal concentrations of carbaryl (5.15, 7.73 and 15.47 microgl-1) for duration of 40 days. The toxic medium was renewed daily. The prawns were fed ad libitum with known energy quantity of boiled goat liver on daily basis. The overall wet weight gain was calculated. The energy lost through unconsumed food (15-60%), faeces (15-109%), ammonia excretion (9-27%) and moults (13-26%) of the prawns were calculated. The feeding rate, the rate and efficiency of absorption, the metabolic and food conversion rates and the gross and net food conversions efficiencies were found to be significantly declined (p<0.05) in test prawns when compared to that of the control. The energy lost through faeces, ammonia excretion and exuvia was found to be significantly elevated (p<0.05) in test prawns than that of the control. The effectof carbaryl on the bioenergetics parameters was severe in the highest sub-lethal concentration, less in the intermediate concentration and least in the lowest sub-lethal concentration. The results indicated that decrease in feeding, absorption, metabolism and food conversion are interdependent and toxicity of carbaryl diverting energy from production to maintenance pathways, which ultimately resulting in declined growth of M. malcolmsonii.     

Beiträge zur Temperaturadaptation des Aales (Anguilla vulgaris)

Previous authors have demonstrated that changes in O₂-consumption of predominant muscle tissue (in vitro) do not account for the variations in O₂-consumption of a resting intact eel induced by differences in adaptation temperature (AT). The present paper is concerned with the metabolism of the eel's lateral muscle. Activity of cytochrome oxidase and succinate dehydrogenase, expressed in terms of O₂-consumption, are higher in the red muscle portion than in the white one (the factors are 6 to 10 and 2 to 3, respectively). In epaxial muscle strips from some 10 cm behind the anus, red muscle tissue comprises between 44% and more than 60% of the total lateral muscle cytochrome oxidase activity, even though, in this region, the relative weight of the red portion makes up only 11 to 14.5% of the total muscle. The red muscle portion is still smaller in the region anterior to the anus, and its total cytochrome oxidase activity (enzyme activity x relative weight) much lower. Cytochrome oxidase activity remains rather unchanged in the white portion of the muscle. Since the metabolism of white muscles is primarily glycolytic, it is remarkable that — with respect to the terminal oxidative metabolism — tissue temperature adaptation in the eel is more pronounced in red muscles than in white ones. In juvenile and adult eels, acclimated to seasonal temperature, the total cytochrome oxidase activity of the white muscle undergoes little change. However, in cold-acclimated adults with extensive pigmentation, aerobic metabolism of red muscles is favoured; this is indicated both by extremely high cytochrome oxidase activity and increased relative muscle weight. Increased aerobic metabolism of red muscle tissue is, therefore, assumed to be important, especially during the winter spawning migration.     

Maturation,fecundity, and spawning strategy of the thornback ray, <Emphasis Type="Italic">Raja clavata</Emphasis>: do reproductive characteristics vary regionally?

In Portuguese waters, thornback ray Raja clavata spawns mainly between May and January, although females and males in spawning condition are found throughout the year. The maturation process can be divided into three main phases by using information on gonad weight, oviducal gland and uterus width in females and on gonad weight, clasper length and sperm duct width in males. Females attain length-at-first-maturity at 784 mm and males at 676 mm, at ages of 7.5 and 5.8 years, respectively. In females larger than length-at-first-maturity, a resting stage was identified characterized by low gonadosomatic index and well-developed oviducal glands and uteri. These results along with the low proportion of adult females that are effectively reproducing per month demonstrated that the thornback ray cannot be considered a continuous spawner as described in other studies. Fecundity was determinate with about 35 eggs released per batch. During the spawning season, a total of four batch episodes occur indicating that the total fecundity was approximately 136 eggs per female. Regional differences may exist in the reproductive strategy of the species, namely on the duration of the spawning season, length-at-first-maturity, and fecundity, which can be related to a more intense fishing pressure in northern European waters.     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

Changes in the biochemical composition of the pyloric caeca of female seastars,Asterias rubens,during their annual reproductive cycle

The biochemical composition of the pyloric caeca of female seastars (Asterias rubens) was studied throughout the annual reproductive cycle. The pyloric caeca index is high during the pseudo-rest stage of the ovaries and decreases gradually during ovarian growth. The pyloric caeca dry weight varies between 20 and 38% of the fresh weight. Lipids, and less prominently carbohydrates (both glycogen and other reducing sugars), seem to constitute the primary nutrient reserves for gonad growth. Proteins are also available to meet ovarian requirements during vitellogenesis. Glycogen levels are high in late autumn and winter and low at spawning time and in summer, whereas levels of the other reducing sugars are high in post-spawning months and subsequently gradually decrease during gonad development. The total lipid level is extremely variable (averaging 75 mg g^-1 organ fresh weight), peak levels of over 200 mg g^-1 being found in some individuals in summer and at spawning time. The free amino acid level is constant at about 20 mg g^-1. The protein level is rather high throughout the annual cycle, and displays only minor changes. The observed patterns of decrease in major constituents suggest a rapid release of nutrient reserves from the pyloric caeca (deposited during the summer) at the onset of vitellogenesis. Moreover, the needs of the ovaries during further development seem to be so tremendous that during the winter months the digested food may be transported to the ovaries without prior deposition in the pyloric caeca. The changes observed in the pyloric caeca are discussed and compared with those occurring simultaneously in the ovaries.     

Reproductive cycles of tropical sea cucumbers (Echinodermata: Holothuroidea) in southern Taiwan

Reproductive cycles of Afrocucumis africana (Dendrochirotida), Actinopyga echinites, Holothuria leucospilota, H. cinerascens, H. difficilis (Aspidochirotida), Synaptamaculata, Opheodesoma grisea, Patinapta taiwaniensis and Polycheira rufescens (Apodida), representing three orders of intertidal holothurians in southern Taiwan, were determined by gonad index and histological examination from March 1990 through September 1991. All nine species of holothurians have annual spawning periods lasting 2 to 4 mo in spring or summer. The early spawning of H. cinerascens and P. rufescens (suspension feeders) suggests that their feeding mode is related to food resources and may influence the reproductive period. In the direct-developing dendrochirotid (suspension feeder), gonad development occurs in cool months, and gametes are released at the beginning of warm months. In indirect-developing aspidochirotid and apodid (deposit feeder), gamete release occurs in late spring or summer and appears to be correlated with summer phytoplankton growth. The holothurians in the present study display spawning periodicity and duration similar to those of temperate species. Four species of holothurians lengthened spawning season with decreased latitude.     

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.