Multiple spawning in the tropical squid Photololigo sp.: what is the cost in somatic growth?

This study assesses the potential of the tropical loliginid squid Photololigo sp. to lay multiple batches of eggs and examines changes in somatic growth during reproduction. Histological analysis of the ovary and the relative size of the oviduct to mantle weight and ovary weight were used to determine the potential for multiple spawning. Ovaries of mature females always had immature and mature oocytes present, suggesting that not all the oocytes were maturing simultaneously and that multiple batches of eggs were being produced. Furthermore, poor correlations of oviduct weight with body size and ovary weight indicated that mature oocytes were not accumulating in the oviduct for a single spawning event. Both these observations supported the hypothesis that Photololigo sp. has the potential to lay multiple batches of eggs throughout its life. Specific growth rates, length-weight relationships, relative growth of somatic and reproductive tissue and microscopic assessment of muscle tissue were compared between immature and mature females. Growth rates of immature females were almost twice as fast as those of mature females. Mature females also had no large muscle fibres present, suggesting that energy for reproduction was mobilised from the muscle tissue.     

Temporal life history plasticity of the Southern Ocean squid <Emphasis Type="Italic">Todarodes filippovae</Emphasis> from waters off Tasmania,Australia

Population dynamics and maturity parameters were analysed for seasonal samples of the oceanic ommastrephid squid Todarodes filippovae from off the coast of Tasmania Australia from 2002 to 2004. Based on assumed daily periodicity in statolith increments, T. filippovae had a life cycle of about a year with the sexually dimorphic females reaching their larger size by predominantly growing faster than males. Due to the small sample size of males, analysis was undertaken on female individuals only. Growth in all samples was best described by a power curve and varied annually, with significantly faster growth in 2001 compared to the subsequent 2 years. Seasonal growth rates also varied with autumn- and winter-hatched squid significantly faster than summer-hatched squid. Spring growth rates were intermediate but not significantly different to the other three seasons. Peak hatching periods occurred in late autumn and early winter. ANOVA revealed a season × year interaction for mantle length and total body weight. Pairwise comparisons showed that the annual differences were likely driven by smaller squid in autumn 2002 compared to autumn samples in the other 2 years. Pairwise comparisons also revealed seasonal differences with winter-caught squid smaller than those from the majority of other samples. There were no seasonal effects on mature female gonad weights but females caught in 2002 had significantly lighter gonad weights than females from the following 2 years. There were no consistent trends among seasons or years in the age structure of mature females. This study revealed the plasticity and flexibility in growth and maturity parameters in this species, with a preference for faster growth during cooler periods.     

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.     

Reproductive biology of the common shrimp Crangon crangon (Decapoda: Crangonidae) in the central Irish Sea

Aspects of the reproductive biology of the common shrimp Crangon crangon (L.) were studied in Port Erin Bay, Isle of Man, Irish Sea. Size at sexual maturity was determined from the proportions of ovigerous females and of females with maturing ovaries. The size at which 50% of females are mature is estimated (±95% confidence intervals) as 12.5±0.48 mm carapace length. Based on the proportions of ovigerous females and of mature females, the main breeding season was from January to June. Mean ovarian dry weights indicated two broods (winter and summer), with females bearing winter broods (WB) having higher gonad indices than those with summer broods (SB). WB females with non-eyed eggs and with eyed eggs differed in the regression of ovarian dry weight on carapace length, indicating preparation for laying a second brood. In both broods the moult stages of berried females were related to egg stage. Moulting will occur following the release of the brood. During embryonic development, mean egg length and egg volume were larger in all stages, and the mean dry weight of individual eggs of all stages heavier, in WB than in SB; there was no difference in egg number, however. Consequently, reproductive investment, the proportion of female weight devoted to egg production, was 67% higher in WB (0.20±0.04) than in SB (0.12±0.03). There is a significant effect of egg volume on brood weight, but not on egg number. In both broods, egg number was a negatively allometric function of female body size in non-eyed eggs and an isometric function of female body size in eyed eggs. Brood mortality during incubation was higher in SB (17%) than WB (10%). Differences in the reproductive variables and investment between the two broods of C. crangon are discussed in the light of reproductive strategies and life history.Communicated by J.P. Thorpe, Port Erin     

Reproduction and condition of female Octopus mimus (Mollusca: Cephalopoda)

The relationship between reproduction and condition was studied in a 15 mo sample of 919 maturing, mature and post-spawning female Octopus mimus (388 to 3714 g) caught in Iquique (North Chile). O. mimus is a semelparous species, with reproduction taking place all year round. Investment in reproductive tissues was, on average, 9.9% of mature female body weight, independent of season. However, somatic growth during maturation was dependent on season and varied between 26 and 63% of the whole-body growth in weight. The condition of females did not vary markedly until spawning, although seasonal variations were apparent, winter being the most unfavourable. Condition deteriorated dramatically after spawning, during parental care of the eggs. During this period, somatic-tissue depletion, mainly from the muscles, was >25% of the total body weight of mature females. The fecundity of O. mimus was probably limited by the costs associated with parental care of the eggs.     

Spatial and temporal variation in growth rates and maturity in the Indo-Pacific squid <Emphasis Type="Italic">Sepioteuthis lessoniana</Emphasis> (Cephalopoda: Loliginidae)

Age, growth and maturity parameters are described for the Indo-Pacific squid Sepioteuthis lessoniana both temporally and spatially (equatorial, tropical and subtropical). Tropical squid that grew through periods of warming water temperatures grew 9% faster than squid that grew through periods of cool water temperatures. The tropical spring-hatched and equatorial squid had similar growth rates (3.24, 3.18 g/day) and these were significantly faster than the tropical summer/autumn hatched squid (2.89 g/day). The oldest squid aged was 224 days, but the majority of individuals were <200 days. Subtropical squid were larger, older and matured later than equatorial and tropical spring-hatched squid. The mean weight of subtropical squid aged between 100 and 150 days was >400 g, 85% greater than tropical and equatorial squid. Geographical differences revealed that subtropical mature winter females and males had mean ages >150 days, respectively 17% and 23% older than their tropical mature winter counterparts. Temporal differences in age at maturity were also evident with tropical winter females and males having a mean age of ~140 days, respectively 41% and 25% older than their summer counterparts. Cooler subtropical and winter tropical squid had the heaviest gonads (>15 g ovaries, >1.5 g testes) compared to summer tropical and equatorial squid. However, relative gonad investment (GSI) values of the cooler squid were significantly lower with cool subtropical and winter tropical females having GSI values <3, which was about half the value of the warmer water females. This study revealed considerable plasticity in the size-at-age of this species. The tropical population had growth parameters that fluctuated between an equatorial strategy (fast growth, small body size, and small gonads) and a subtropical strategy (large body size, slower growth, and large gonads) depending on season.     

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.     

Flexible reproductive strategies in the squid Loligo forbesi

Body size at sexual maturity, egg-size distributions, and potential reproductive output have been estimated for female and male squid, Loligo forbesi Steenstrup, off the west coast of Scotland. Two size modes at maturity were found in both sexes, but separation into size cohorts was more pronounced in males (180 and 350 mm mantle length, ML). Preliminary ageing studies based on statolith ring-counts suggest that these size modes are not due to different age groups at breeding. Females have a single size mode of mature eggs in the proximal oviduct, but may have at least two size modes of eggs within the ovary. This finding is interpreted as evidence of batch-spawning in this squid. There was a weak relationship between total egg numbers (range 1000 to 16000) and body size (range 196 to 318 mm ML) and between mature egg size and body size. Males showed a strong positive relationship between spermatophore length and body length but a weak relationship between total number of spermatophores and body size. The results are discussed in the context of flexibility of breeding strategies in the loliginids and variety of life-cycle patterns.     

Evidence for multiple spawning in the tropical oceanic squid Stenoteuthis oualaniensis (Teuthoidea: Ommastrephidae)

Indirect evidence suggests that Stenoteuthis oualaniensis is a multiple spawner, a reproductive strategy not reported previously for squid. After spawning once, S. oualaniensis apparently continues to feed, grow, and mature additional oocytes before spawning again. Squid were caught by jigging at night in waters around the main Hawaiian Islands during four cruises from 1981 to 1987. All females from 200 mm mantle length (ML) to at least 318 mm ML were sexually mature, indicating that considerable growth takes place after reaching sexual maturity. Size-frequency analyses of ovarian oocytes revealed continuous egg production throughout adult life. Mature ova are stored in the oviducts, but oviduct fullness is unrelated to body size, indicating repeat spawning.     

Reproductive cycle and maternal effects on offspring size and number in the neogastropod Buccinum undatum (L.)

This paper examines relationships between maternal phenotype and offspring phenotype in the commercially exploited marine neogastropod Buccinum undatum (L.). The study also aimed to estimate the effects of female size on subsequent recruitment, and to determine the reproductive cycle of B. undatum in Swedish waters. The evaluation of gonad development in field-sampled animals from August 1998 to February 2000 revealed a definite annual reproductive cycle with a single major egg-laying period in the autumn. In addition, adult whelks of varying sizes from two locations and in two consecutive years were kept in aquaria to study the influence of female size on fecundity. Twenty-seven females deposited eggs in the laboratory. Offspring quality, measured as egg and hatchling size, was independent of female size, although larger females produced more eggs and hatchlings than smaller females, and thus probably account for a disproportionately large share of potential recruitment; the relative fecundity (number of eggs or hatchlings per gram of body weight) was constant with female size. I found no evidence for a longer time interval between successive egg-layings in larger females, i.e. mature females of all sizes probably spawn equally often. The present results therefore indicate that spawning-stock biomass is probably a better measure of spawning-stock size than the number of mature individuals in the population. Effects of an individual female's characteristics on stock productivity have largely been overlooked by fishery managers but could help to explain the weakness often found in stock-recruitment relationships.     

Reproductive development of the barnacle Chthamalus malayensis in Hong Kong: implications for the life-history patterns of barnacles on seasonal, tropical shores

Hong Kong, lying just below the Tropic of Cancer (22°17′N, 114°09′E), experiences a strongly seasonal environment, with a cool almost temperate winter and a hot, tropical, summer. Histological sectioning of the gonads of the high-shore barnacle, Chthamalus malayensis Pilsbry, showed a seasonal trend in the development of its reproductive organs. Four stages of female gonad development were identified according to the cell types present: post-spawning, resting, growth and mature stages. The female gonad was mature from April to November, which was related to seawater temperatures, and entered a resting phase from December to March. Although the male gonad showed a seasonal developmental trend and reached maximum maturity in summer, the seminal vesicles were full of spermatozoa and functional throughout the year. The reproductive season of this species is therefore solely dependant upon the maturity of the female gonad. The estimated maximum number of broods per year was up to 10 and the maximum number of eggs produced per brood can reach 3,000 eggs. The minimum size for female gonad maturity was 6 mm rostro-carinal diameter (RCD) at which size, the barnacles were ~6-month old. Sperm production occurred at a smaller size (2 mm=2-month old). Compared with Chthamalus montagui and Chthamalus stellatus from temperate regions, C. malayensis produced a greater number of broods per year, had a longer reproductive period and faster gonad development. Chthamaloid barnacles in tropical regions may, therefore, invest more energy per year in reproduction during their life span. Contrary to the seasonal gonad developmental pattern of C. malayensis in the present study, however, C. malayensis in Singapore (which experiences only slight seasonal variation) had mature female and male gonads throughout the year, further supporting the strong role of climatic conditions effecting the reproductive biology of barnacles.     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Geographic variation in maternal investment: acidity affects egg size and fecundity in Rana arvalis

Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.     

Female benefit,male risk: Polyandry in the true armyworm Pseudaletia unipuncta

In Lepidoptera polyandry is common and females may increase their lifetime reproductive output through repeated matings if they acquire essential resources from male ejaculates. However, the paternity of males mating with previously-mated females is far from assured unless sperm precedence is absolute. In this study on the polyandrous armyworm, Pseudaletia unipuncta, we used two strains of male (the black-eyed wild type and a red-eyed homozygous, recessive mutant), mated with red-eyed females, to determine (i) whether male investment has any impact on female reproductive output, and (ii) if females do benefit from multiple matings, to what extent males fertilize the eggs to which they contributed. Multiple mating resulted in a significant increase in both the fecundity and longevity of females. However, the degree of sperm precedence (those eggs fertilized by the second male) varied from 0–100%, but was not affected by either male size or age, or by the duration of copulation. In cases where sperm precedence was <50% (x = 12%) females produced significantly more eggs (1384 versus 940) prior to the second mating than females where sperm precedence was >50% (x = 89%), indicating that the quality of the first mating influenced the fertilization success of the female's second mate.     

Fecundity,egg weight and longevity in relation to multiple matings in females of the monarch butterfly

Summary In many insects nutrients transferred by the male at mating are later incorporated into both the eggs and soma of the mated females. Accordingly, it has been suggested that insect females can use these male-derived nutrients for somatic maintenance and enhancement of their fecundity and fitness of their offspring. I this paper we tested the validity of these predictions by studying the longevity and reproductive output of 1, 2, 3, 4 or 5 times mated female monarchs Danaus plexippus, a nectar-feeding butterfly that is long-lived and strongly polyandrous and emerges as an adult without mature eggs. Females mated five times received ejaculates that corresponded to an average of 38% of their body weight at eclosion. However, we found that the number of times females had mated had no effect on their longevity, life-time fecundity, or egg weight. Although negative evidence should always be interpreted with caution, our study suggests that male-derived nutrients are less important for female longevity and reproductive output than are larval and adult food.     

Support for maternal manipulation of developmental nutrition in a facultatively eusocial bee, <Emphasis Type="Italic">Megalopta genalis</Emphasis> (Halictidae)

Developmental maternal effects are a potentially important source of phenotypic variation, but they can be difficult to distinguish from other environmental factors. This is an important distinction within the context of social evolution, because if variation in offspring helping behavior is due to maternal manipulation, social selection may act on maternal phenotypes, as well as those of offspring. Factors correlated with social castes have been linked to variation in developmental nutrition, which might provide opportunity for females to manipulate the social behavior of their offspring. Megalopta genalis is a mass-provisioning facultatively eusocial sweat bee for which production of males and females in social and solitary nests is concurrent and asynchronous. Female offspring may become either gynes (reproductive dispersers) or workers (non-reproductive helpers). We predicted that if maternal manipulation plays a role in M. genalis caste determination, investment in daughters should vary more than for sons. The mass and protein content of pollen stores provided to female offspring varied significantly more than those of males, but volume and sugar content did not. Sugar content varied more among female eggs in social nests than in solitary nests. Provisions were larger, with higher nutrient content, for female eggs and in social nests. Adult females and males show different patterns of allometry, and their investment ratio ranged from 1.23 to 1.69. Adult body weight varied more for females than males, possibly reflecting increased variation in maternal investment in female offspring. These differences are consistent with a role for maternal manipulation in the social plasticity observed in M. genalis.     

No cost of male mating experience on female reproductive success in the almond moth, <Emphasis Type="BoldItalic">Cadra cautella</Emphasis> (Lepidoptera; Pyralidae)

Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness advantage to females.     

Strontium/Calcium ratios in statoliths of the neon flying squid, Ommastrephes bartrami (Cephalopoda), in the North Pacific Ocean

Strontium to calcium ratios were observed along longitudinal sections of statoliths of nine neon flying squid, Ommastrephes bartrami (LeSueur, 1821), including three mature females (422 to 454 mm mantle length, ML; 207 to 306 d old) obtained from the North Pacific (27–35°N; 144–150°E) during winter and six immature males and females (187 to 226 mm ML; 126 to 164 d old) collected from 39°N; 145°E and 39°N; 169°W during summer. The distances between the nucleus (core) and the edge of the dorsal dome were approximately 660 to 690 μm in mature females and 450 to 510 μm in the immature squid. Sr/Ca ratios were determined at intervals of 30 μm between the nucleus and edge of the dorsal dome. Sr/Ca ratios were higher in areas near the nuclei and peripheral portions of the dorsal dome than in the middle portions of the statoliths (270 to 420 μm from the nuclei, corresponding to ages of 60 to 90 d) in mature females; thus a U-shaped pattern was evident. Sr/Ca ratios in the six immature squid decreased from nucleus to the dorsal dome; in three squid the ratios slightly increased toward the dorsal dome edge. The observed Sr/Ca ratios in immature squid were considered to represent younger portions of the U-shaped pattern. In the present study we discuss this pattern in relation to environmental and biological conditions of O. bartrami, which undertakes seasonal migrations between spawning grounds in the Subtropical Domain and feeding grounds in the Subarctic Domain and Transitional Zone in the North Pacific Ocean. Although Sr/Ca ratios are potentially affected by ambient water temperature and ontogenetic conditions, including somatic growth and statolith growth, it was impossible to evaluate each environmental and biological effect separately, as variations in these factors are complicated and effects could be interdependent. Received: 11 April 1997 / Accepted: 27 December 1997     

Reproduction and growth of the freshwater prawn, Palaemon paucidens (Decapoda: Palaemonidae) in a lake of Korea

This study analyzed the reproduction and growth of freshwater prawn, Palaemon paucidens De Haan 1844 from Suk-dang lake(Korea). The analysis of the sex ratio showed a higher proportion of males than that of females. The average size of the eggs was 6.12 (+/-0.55) mm3 in the non-eyed stage (stageA) and 7.20 (+/-0.86)mm3 in the eyed stage (stage B). The reproductive output (RO) calculated with the dried-body weight of an incubating female and the weight of the eggs in dried condition was 26.97% (n = 17) of the average females weight. Incubating prawn appeared in April, and the gonadosomatic indices(GSI) showed the highest value during three months from January to March. Based on the month when there was a high proportion of incubating females with a high GSI, the estimated spawning season of P. paucidens was April. The maturity of the female was evaluated by the development of the ovaries and the existence of eggs. The average body length when 50% of the females in the group reached maturity was 8.55 (+/-2.74) mm. The analysis of the length-frequency distribution showed that the life span of P. paucidens ranged from 12 to 13 months. Females mature faster than males.     

Lipids in the digestive gland and the gonad of immature and mature Sepia officinalis (Mollusca: Cephalopoda)

The lipid content of the digestive gland and the gonad of cuttlefishes (Sepia officinalis L.) caught off the French coasts in 1979–1980 was determined and analysed by thinlayer chromatography. The total sterol and phospholipid levels were also determined. The results from immature and mature females and males were compared in an attempt to correlate variations of the lipids with sex or state of gonad maturation. The fluctuations in the lipid level of the digestive gland, which acts as a storage organ, seemed far better related to the diet than to sex or sexual maturity. In the gonads, the lipids were few and seemed to be mostly membrane constituents. However, the increase of phospholipids in the mature ovary could be related to yolk synthesis.