Experimental induction of gonadal maturation and spawning in the giant jellyfish <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> (Scyphozoa: Rhizostomeae)

We studied the processes of gonadal maturation, spawning, fertilization and embryonic development of the giant jellyfish Nemopilema nomurai, which has been blooming in recent years in the Sea of Japan. Healthy medusae actively swimming offshore had immature gonads, but damaged and ashore-drifted animals had relatively mature gonads. The animals maintained in a small net on the nearshore to mimic the drifted condition showed induced gonadal maturation by the increase in oocyte diameter and darkness in the matrix. A similar maturation process also occurred in isolated pieces of ovary incubated at the same temperature. Fully grown oocytes that probably stopped at the prophase of the first meiosis reinitiated maturation divisions in response to light exposure, and massive spawnings occurred 80–100 min after the exposure. The spawned oocytes were first released within the subgenital sinus, then transported peripherally and finally shed into the gastrovascular cavity. Maturation and spawning in male gonads were similar to those of females except that the male spawning occurred within 30 min after the light exposure and always preceded the female spawning. Hence, it was suggested that fertilization might take place in the female gastrovascular cavity by the sperm that came in from surrounding seawater by animals’ pumping activities after dawn. Having mechanical damages might promote fertilization success of this species in the Sea of Japan.     

Reproductive cycles of three dominant Calanus species in Disko Bay, West Greenland

This study describes the annual reproductive cycles of the three dominant Calanus species, C. finmarchicus, C. glacialis and C. hyperboreus, in Disko Bay (West Greenland) in relation to seasonal phytoplankton development. Relative abundance of females, copepodite stage V (CV) and males, and the developmental stage of the female gonad were examined from plankton samples collected at weekly to monthly intervals from May 1996 to June 1997 with a WP2 net or a pump. During spring and summer, egg production rates were determined. Females of all three species were present year round. Maximum relative abundance was reached by C. hyperboreus females at the beginning of February, by C. glacialis in mid-February, and by C. finmarchicus in April. All three species reproduced successfully in Disko Bay. Their reproductive cycles were considerably different with respect to the timing of final gonad maturation and spawning, and hence in their relation to seasonal phytoplankton development. In all three species, early gonad development took place during winter, before living food became plentiful, suggesting that these processes were largely food independent. Final gonad maturation and spawning in C. finmarchicus was related to the phytoplankton concentration, reflecting that final gonad maturation processes are food dependent in this species. C. glacialis females matured and spawned prior to the spring bloom. Our results indicate that first internal lipid stores and later ice alga grazing supplied final gonad maturation and egg production. Maximum egg production rates of C. glacialis were found in spring and summer, when the chlorophyll a concentration was high. Mature female C. hyperboreus were found from February until mid-April, when the chlorophyll a concentration was still low. In this species, reproductive activity was decoupled from phytoplankton development, and final maturation processes and spawning were solely fuelled by internal energy stores.     

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.     

Size-related reproductive biology of the infaunal bivalve<Emphasis Type="Italic"> Abra alba</Emphasis> (Bivalvia) in Kinsale Harbour (south coast of Ireland)

The reproductive behaviour of 248 specimens comprising four size classes of the infaunal bivalve Abra alba (Wood) was investigated between September 1994 and November 1995 in Kinsale Harbour. Small males returned a very small proportion of sexually mature and spawning animals. By contrast, no spawning females were found in this size class, suggesting that males develop and mature at a smaller size than females. Mean gonad maturity indices indicated that, except for the smallest size class, A. alba exhibits two spawning events in 1 year, i.e. a lesser spring and a major autumn spawning event. The spring event was well synchronised between males and females, while the autumn spawning was more broadly synchronised between the sexes. Only one spawning event was recorded for very large males and females, also occurring in spring. Spawning stages were recorded for both sexes over all size classes between February and November, indicating only a short period of sexual repose, if any. Oocyte diameters did not show a discrete size range from developing to spawning stages. All stages were seen to contain varying proportions of smaller, less mature oocytes, suggesting that large A. alba females do not develop and spawn en bloc, but release their gametes periodically. Maturity indices showed only a broad similarity with off-bottom temperatures.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.     

Energy investment in the annual reproduction cycle of female red porgy, <Emphasis Type="Italic">Pagrus pagrus</Emphasis> (L.)

Seasonal cycle in energy content of gonad, perivisceral fat, liver, and muscle was used to estimate the energy cost of spawning of female red porgy (Pagrus pagrus) from Buenos Aires coasts, Argentina. Spawning was observed primarily between November and January, although a low-intensity spawning was also observed at the beginning of this period. The energy required for gonad development of female red porgy (the difference between the maximum and minimum energy content of ovaries along the year) was 2,652 J g⁻¹ dry weight or 558 kJ. Female red porgy expend 22.8% of their total body energy (3,481 kJ) during the spawning period. Of this energy, relative contributions from different sources were as follows: ovaries 62.5% (496 kJ), perivisceral fat 17.8% (142 kJ), liver 10.3% (82 kJ), and muscles 9.4% (75 kJ). Female spawners lost about 7.2% (119 g) of their total body weight (1,654 g) in the spawning period. From the 7.2% of the total weight lost, female gonad represented a relative reduction of 73.8% (87.6 g), the perivisceral fat declined by 5.7% (6.8 g), the liver declined by 10.6% (12.5 g), and muscle tissue exhibited a 10% loss in total weight (11.9 g). Reproductive effort represented 7.6% of food energy, while resting metabolism accounted for almost 38% of food energy. Despite the fact that red porgy females feed throughout the entire spawning period, it is shown that between pre-spawning (September, water temperature 15°C) and mid-spawning (December, water temperature 19°C) they lost 45 and 38% of their muscle and liver lipids, respectively, and 89% of the perivisceral fat associated with the gut and body cavity. Red porgy used both their energy reserves and food energy during ovarian maturation. This asynchronous species have a strategy of a lower energy allocation and a higher dependence of food during the spawning time (late-spring and summer) for successful reproduction.     

Stereological methods applied to reproductive cycle of Tapes rhomboides

A population of Tapes rhomboides (Pennant) in the Bay of St. Malo, France, was studied for one and a half years (July 1984–October 1985) to determine spawning frequency and fecundity under natural conditions, using the techniques of qualitative histological staging, condition index calculation and quantitative stereology. Spawning took place twice a year, in late May and in July/September. There was an extended winter resting period. Gonad development and oocyte production were positively correlated with female body size. The annual fecundity in a 40 mm clam was about 5x10⁵. Stereological techniques provided accurate information on oocyte dynamics within a gonad and the relationship with atretic processes. Each oocyte generation from the onset of the cycle to the winter resting stage was spawned eventually if conditions were suitable, or the oocytes were resorbed. Atresia of oocytes was seen throughout the reproductive period, but especially after the spring spawning and at the end of the summer.     

Reproduction of the commercial sea cucumber Holothuria scabra (Echinodermata: Holothuroidea) in the Solomon Islands

Over a 3-year period (1996-1998), reproduction of the commercial sea cucumber Holothuria scabra (Jaeger, 1833) was investigated in the Solomon Islands to determine the spawning pattern and whether gametogenesis is continuous or seasonal. The gonad consisted of a single cohort of tubules that developed uniformly. Macroscopic examination of the gonads revealed that mature gametes were present throughout the year. Individuals with gonads at different stages of maturity were present in most samples. Partly spawned gonads were prevalent in females, whereas mature gonads were prevalent in males. The time at which the peak gonad index was recorded differed among years. Although gametogenesis was continuous, with a potential for prolonged gamete release, a period of enhanced spawning occurred during the dry season, from September to December. Maximum gonad indices were reached prior to and during this period of enhanced spawning. Histology revealed that gametogenesis reinitiated in partly spawned gonads, resulting in the presence of gametes at different stages of development in the gonad. The uniform growth of gonad tubules indicated that H. scabra does not conform to the progressive tubule recruitment model described for holothurian oogenesis. Continuous reproduction in H. scabra and prolonged availability of mature gametes would facilitate use of this species for aquaculture.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.     

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     

Development and expression of sexual maturity in femaleCalanus pacificus (Copepoda: Calanoida) in relation to food quality

Dinoflagellates, which comprise an important part of the phytoplankton in the neritic region of the Southern California Bight, are known to be grazed byCalanus pacificus; rates of ingestion, development and survival of nauplii are influenced by the food quality of dinoflagellates. We have examined the effects of dinoflagellate food quality on reproduction ofC. pacificus females sampled in La Jolly Bay (32°N; 117°W) between 6 April and 4 May 1988. Four sets of experiments were conducted in which copepods were fed five different species:Gymnodinium splendens, Gyrodinium resplendens., G. dorsum, Exuviella mariae lebourae andGymnodinium simplex. These species were selected on the criterion of their success in supporting the growth of nauplii in previous rearing experiments. The experimental culture concentration was maintained at 100µg C 1^–1, near that of the natural seston. As indices of successful reproduction, we measured the percentage of spawning females and the number of eggs laid per female per day. Laboratory results were compared to stages of ovarian maturation of wild females. Four stages of maturation were determined using histological and microscopic observations.Gymnodinium splendens, Exuviella mariae lebourae, andG. simplex yielded clearly inferior results: few females were able to lay eggs repetitively over the first few days of confinement.Gyrodinium resplendens and, above all,G. dorsum yielded the best results. The stage of sexual maturation in situ appears to influence the rhythmicity of egg-laying events. In the presence of high-quality food, females are able to maintain high reproductive rates by sustaining the maturation of successive batches of eggs; good food quality appears to assure this secondary vitellogenesis and the rapid turnover of ovocytes.     

Temporal shifts in the allocation of energy in the arrow squid,<Emphasis Type="Italic"> Nototodarus gouldi</Emphasis>: sex-specific responses

Squid typically display considerable intra-specific plasticity in size and age-at-maturity in response to ambient environmental conditions, yet little is known of the mechanisms driving these variations. We examined the intra-specific variability in Nototodarus gouldi reproductive traits to determine patterns of energy allocation between somatic and reproductive processes over short temporal scales. Females caught during the cool months of May and July were larger, had slower lifetime growth, lower gonad investment, and better somatic condition than females caught during the warmer months, suggesting a trade-off between gonad investment and somatic condition in females. On the other hand, males showed a tight coupling between somatic condition and gonad investment for most months, with increases in somatic and gonad tissue occurring concurrently. In male squid, an increase in lifetime growth rate was coupled with an increase in the relative weights of somatic and reproductive structures, whereas in females, percent increase in body weight per day was correlated only with gonad development. Patterns of repro-somatic investment in mature females had implications for spawning strategies, since female squid with higher levels of gonad investment apparently released batches of eggs together as a group, regardless of body size, whereas females with low gonad investment possibly spawned their eggs independently of one another. In terms of life-history theory, male squid were able to respond rapidly to environmental fluctuations without compromising either the gonad or the soma. However, although mature females did not appear to respond as quickly to ambient conditions, female squid possibly produced two different reproductive strategies, possibly to maximise offspring survival in either a stable or a variable environment. It seems from our study that monthly variations in ambient conditions may have large effects on life-history strategies.Communicated by M.S. Johnson, Crawley     

Multi-year observations on the gametogenic ecology of the Antarctic seastar <Emphasis Type="Italic">Odontaster validus</Emphasis>

This study reports the first multi-year observations on the reproductive patterns for an Antarctic predator/scavenger, Odontaster validus (Koehler 1912). Seastars were collected monthly from a shallow site (15–20 m depth) near the British Antarctic Survey (BAS) Rothera Research Station (Adelaide Island, 67°34′S 68°08′W) from July 1997 to January 2001. Reproductive condition, oocyte size frequencies and spermatogenesis were examined in at least ten seastars each month using histological and image analysis techniques. Gonad indices (GI) and pyloric caeca indices (PI) were also examined in the same samples. Female and male GIs varied seasonally, in parallel with a reduction in the proportion of large oocytes and mature sperm in the gonad in August to mid-October following winter spawning. Despite there being remarkable consistency in the timing of spawning from year to year, differences in the reproductive condition of individuals were apparent. Patterns in the digestive tissues also varied with season, peaking in December and reaching a minimum in February in two of the three study years. This weaker annual pattern may partly reflect the varied diet of this predator/scavenger species, which is not directly dependant on the timing and magnitude of the annual phytoplankton bloom. Pooled oocyte size distributions and residual analysis suggested that oogenesis progressed over 18–24 months, with the largest of the two size classes (maximum diameter = 183 μm) being spawned annually. This pattern of oocyte growth and spawning was previously reported in the early 1960s for an O. validus population from McMurdo Sound, which lies south of Rothera by 10° latitude. The extremely catholic diet of this predator/scavenger suggests the reproductive patterns of the seastar will be less susceptible to changes in food supply compared to polar suspension feeders or deposit feeders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Growth and reproductive biology of the small penaeid shrimp Trachysalambria curvirostris in Tokyo Bay

We investigated the growth and reproductive biology of the penaeid shrimp Trachysalambria curvirostris in Tokyo Bay, Japan, by monthly bottom-trawl surveys from May 2002 to December 2004. We also examined oogenesis in T. curvirostris by histological observation of the ovary. Females grew faster and attained a larger body size for age than males. The growth rate was high in summer and low in winter and was likely to be associated with seasonal changes in water temperature. The carapace length (CL) at which 50% of females contained vitellogenic oocytes was estimated to be 17.0 mm. The reproductive season extended from May to October. Young-of-the-year appeared in October and could be traced across the months on CL histograms to the following September or October, indicating a 1-year life cycle. This extended reproductive season, together with our observation of asynchronous development of oocytes in the ovary, suggests that multiple spawning by individual females may occur during the reproductive season. Postovulated oocytes were not found among the samples we collected during the daytime, suggesting that final oocyte maturation and spawning occur at night. Cortical crypts in the cytoplasm of the oocyte, considered to be a general feature of oogenesis in penaeid shrimps, were not found in T. curvirostris, even in oocytes undergoing germinal vesicle breakdown. This result implies that the cortical reaction after spawning of T. curvirostris may be different from that of other penaeid shrimps.     

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     

Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Egg production in the arrow squid Nototodarus gouldi (Cephalopoda: Ommastrephidae), fast and furious or slow and steady?

The process of reproductive maturation and egg release was examined in the temperate shelf squid Nototodarus gouldi. The energy allocation between somatic and reproductive growth from juvenile to mature adult was investigated throughout the life span to determine the underlying energetic strategy adopted by individuals. The relative weight of the mantle, fin and digestive gland remained unchanged during ovarian development, with no significant correlations found between the mantle length (ML)-gonad residuals and the ML-mantle (r=0.01, P>0.05), ML-fin (r=0.07, P>0.05) and ML-digestive gland (r=0.07, P>0.05) residuals. This suggested that energy was not being diverted away from somatic growth during sexual development, and consequently neither muscle nor digestive gland was being utilised as an energy store. Since squid in all maturity stages contained some food in their stomachs (e.g. 66.7% of mature animals), it is likely that the cost of maturation in this species is largely being met by food intake. The energy investment in reproductive tissues was relatively low (mean gonado-somatic index for mature individuals was 9.29% - 0.40%), indicating that only small amounts of energy were being allocated to reproduction at anyone point in time, which is characteristic of a multiple-spawning strategy. Furthermore, oviduct weight was not correlated with body size (r=0.256, P>0.05), suggesting that eggs are not stored for a single release. In all except one individual, ovary weight was consistently heavier than oviduct weight, suggesting that the ovary is not being depleted of oocytes as mature ova move into the oviducts. Additionally, the ovaries of mature females contained a range of oocyte sizes with discrete peaks, indicating a continued production and development of oocyte cohorts. The presence of some individuals with stretched empty oviducts is further evidence that the reproductive strategy of N. gouldi is slow and steady, with eggs possibly being released in discrete batches over a period of time.