Timecourse of oocyte development in saithe Pollachius virens

Wild caught North Sea saithe Pollachius virens were monitored for growth, sex steroid profiles and oocyte development pre‐spawning and measured for egg size and group fecundity during the spawning season in the laboratory. Vitellogenesis commenced in late October–early November, at a leading cohort size (C_L) of c. 250 µm, after which oocytes grew rapidly in size until spawning started in February. Notably, a distinct cortical alveoli stage was virtually absent with yolk granules observed in developing oocytes at the very beginning of vitellogenesis. Little atresia was observed pre‐spawning, but atretic re‐absorption of remnant oocytes containing yolk granules was found in all females immediately post‐spawning. As expected, concentrations of sex steroids, oestradiol‐17β (females), testosterone (both sexes) and 11‐ketotestosterone (both sexes), increased pre‐spawning before dropping post‐spawning. The present experiment provides the first validation of sex steroid levels in P. virens. Post‐ovulatory follicles were visible in histological sections from female gonads 9–11 months post‐spawning, but then disappeared. Spawning commenced around a C_L of c. 750 µm (700–800 µm). Hydrated oocytes (eggs) measured between 1·04 and 1·31 mm (mean = 1·18 mm) with decreasing sizes towards the end of spawning. The average estimated realized fecundity was c. 0·84 million eggs (median female total length, L_T = 60 cm). Spawning lasted from 13 February to 29 March.     

Seasonal prevalence and intensity of follicular atresia in Baltic cod Gadus morhua callarias L.

In the present study, 307 ovaries of eastern Baltic cod Gadus morhua callarias sampled during the prespawning and spawning season 2000 were analysed histologically to estimate the seasonal prevalence and intensity of atresia. The number of atretic oocytes per ovary was estimated using a combination of the physical disector method and volume fraction (Delesse principle). Atretic oocytes were observed in 32% of the ovaries. Prevalence of atresia was independent of female size, but increased significantly with declining female condition from prespawning and through the spawning stages. The relative intensity of atresia, i.e. number of atretic oocytes in relation to normally developed vitellogenic oocytes, was low amounting to 1·4% on average. Similar to prevalence, relative intensity of atresia differed significantly between maturity stages and increased with decreasing female condition. The population egg loss due to atresia amounted to 4·6% indicating that Baltic cod was performing close to maximum productivity, i.e. potential egg production.     

Ovarian dynamics and fecundity regulation in blueback herring,Alosa aestivalis,from the Connecticut River,US

We analyzed ovarian dynamics of anadromous blueback herring, Alosa aestivalis, in Connecticut River with the principal aim of exploring oocyte recruitment and how it shapes the fecundity pattern. We examined the oocyte release strategy and analyzed spawning cyclicity by linking oocyte growth to the degeneration of postovulatory follicles. Females were accordingly classified as pre-spawners, early and late active spawners, and oocyte recruitment intensity was compared among the different spawning phases. Oocyte recruitment occurred continuously and in parallel with spawning activity, a pattern which is diagnostic of indeterminate fecundity. However, both fecundity and oocyte recruitment intensity progressively decreased (tapered) throughout spawning, until the ovary was depleted of vitellogenic oocytes. There was no massive atresia of vitellogenic oocytes at the end of the spawning season, which is atypical of indeterminate spawners. We propose that tapering in oocyte recruitment and fecundity is an adaptation to the high energetic expenditure of the upstream spawning migration.     

Primary and secondary oocyte growth dynamics in anadromous semelparous Allis shad Alosa alosa

We analysed the ovarian dynamics of the anadromous semelparous allis shad Alosa alosa for which our working hypothesis was that mature pre-spawning females would have very low or even exhausted primary growth (PG) oocyte reserves; semelparity has been linked with the depletion of the pool of PG oocytes. To test this hypothesis, the PG oocytes were enumerated, their recruitment pattern to the secondary growth (SG) phase was analysed and their potential replenishment from the pool of oogonia was examined in females caught very close to the Mondego River mouth, in central Portugal and along the river. The development of the SG oocytes was also analysed, the fecundity (batch, total and annual) values were estimated and the intensity of atresia was quantified. Ovarian samples and histological sections were investigated in parallel. A dynamic recruitment pattern of PG oocytes to the SG phase was revealed, where all PG oocytes were recruited and were not replenished by oogonia. Annual fecundity was subject to down-regulation due to atresia prior to spawning and its size was multiple times higher than the size of batch fecundity. Lack of population synchronicity in ovarian development and spawning migration was also observed. This multifaceted analysis of the ovarian dynamics of this species will contribute to management efforts for this critically endangered and economically important fish throughout its geographical distribution. The results reported in this study will also assist in unravelling the complexity of the early processes of oogenesis in fish.     

The reproductive cycle of female Ballan wrasse Labrus bergylta in high latitude,temperate waters

This 2 year study examined the reproductive cycle of wild female Ballan wrasse Labrus bergylta in western Norway as a precursor to captive breeding trials. Light microscopy of ovarian histology was used to stage gonad maturity and enzyme‐linked immuno‐absorbent assay (ELISA) to measure plasma concentrations of the sex steroids testosterone (T) and 17β‐oestradiol (E₂). Ovarian recrudescence began in late autumn to early winter with the growth of previtellogenic oocytes and the formation of cortical alveoli. Vitellogenic oocytes developed from January to June and ovaries containing postovulatory follicles (POF) were present between May and June. These POF occurred simultaneously among other late maturity stage oocytes. Plasma steroid concentration and organo‐somatic indices increased over winter and spring. Maximal (mean ±s.e .) values of plasma T (0·95 ± 0·26 ng ml^?1), E₂ (1·75 ± 0·43 ng ml^?1) and gonado‐somatic index (I_G; 10·71 ± 0·81) occurred in April and May and decreased greatly in July when only postspawned fish with atretic ovaries occurred. Evidence indicates that L. bergylta are group‐synchronous multiple spawners with spawning occurring in spring and peaking in May. A short resting period may occur between late summer and autumn when previtellogenic oocytes predominate and steroid levels are minimal.     

Reproductive strategy and spawning activity of the European hake Merluccius merluccius (L.) in the Bay of Biscay

Histological examination of ovaries from European hake Merluccius merluccius from the Bay of Biscay sampled on a monthly basis from December 1996 to October 1997 demonstrated that European hake exhibit indeterminate fecundity. Oocyte development is asynchronous with a continuous oocyte size-frequency distribution in prespawning, spawning and postspawning females, and a gradual decrease in the mean diameter of advanced yolked oocytes as spawning proceeds. Spawning European hake were observed throughout the 11 month study period. The spawning fraction ranged from 0·085 to 0·207, which is equivalent to a batch interval of 5–12 days. The spawning fraction and proportion of spawning females were highest from January to March. Seasonal variation of atresia was inversely related to spawning fraction. Extensive atresia in individual fish was observed in September and October, when the spawning fraction was low.     

Reproductive Biology of Three Commercially Important Hemiramphid Species in South-eastern Australia

Synopsis The reproductive biology of eastern sea garfish Hyporhamphus australis, eastern river garfish H. regularis ardelio, and snub-nosed garfish Arrhamphus sclerolepis were described throughout their respective ranges in the coastal waters of New South Wales (NSW), Australia. Peaks in gonadosomatic indices indicated that spawning of eastern sea garfish occurred in late spring and early summer (November–December) on the south coast of NSW, and in winter and early spring (June–September) on the north coast. Eastern river garfish spawned between July and December in NSW estuaries and snub-nosed garfish spawned between October and January in the Clarence River. The sex ratios in commercial catches of eastern sea garfish from the north coast of NSW were biased toward male fish, but approached equality for fish caught from the south coast. Sex ratios were significantly biased toward female snub-nosed garfish, and female eastern river garfish from all estuaries sampled. Mean (±SE) batch fecundity was 1498 ± 110 (range: 98 – 3449) ripe oocytes per female for eastern sea garfish, 917 ± 36 (range: 102 – 2268) ripe oocytes per female for eastern river garfish, and 716 ± 104 (range: 20 – 2956) ripe oocytes per female for snub-nosed garfish across the range of mature sizes examined. There was a linear relationship between batch fecundity and fish size for all three species of garfish. Eastern sea garfish approached 50% maturity at a larger size than snub-nosed, or eastern river garfish. Size at 50% maturity was similar for male and female eastern river and snub-nosed garfish, but male eastern sea garfish matured at a significantly smaller size than females. All three species appear capable of spawning in the spawning season immediately following the one in which they were born. Mature female fish of all three species had distributions of oocyte diameters consisting of three or four modes, which strongly suggests asynchronous oocyte development and a multiple spawning strategy during the spawning season. Implications for the management of garfish fisheries in NSW are also discussed.     

Reproductive patterns of blacktail comber (Serranus atricauda,Serranidae) from south-west Portugal seamounts

The knowledge of the reproductive strategy is essential for the comprehensive understanding of the population dynamics of any fish species. This study intends to define the spawning season, verify the type of fecundity regulation and estimate relative annual fecundity for Serranus atricauda inhabiting the south-west of Portugal seamounts (Gorringe, Ampère and Lion). Monthly samples, between September 2010 and December 2011, were obtained from commercial vessels. This species is synchronously hermaphroditic, and mature individuals were found all year round, with gonadosomatic index peak from February to September. No clear relation between hepatosomatic index and Fulton’s condition factor and gonadal development was found, and therefore, it is probable that reproductive success depends more on environmental conditions and food availability during the spawning season than on body energy reserves. No gap exists between pre-vitellogenic and vitellogenic oocytes and the oocyte size-frequency distribution during the main spawning season show no dominant cohort progressing through time. The number of standing stock of advanced vitellogenic oocytes showed an increase trend along the spawning season decreasing only at its end and the relative intensity of α-atresia was always low throughout this period with an increase towards the end. All these criteria corroborate the indeterminate fecundity type assigned to S. atricauda in previous studies. Mean relative annual fecundity estimated was 5,655 ± 4,321 oocytes per gram of gutted weight.     

Reproductive biology of albacore Thunnus alalunga

Reproductive variables in albacore Thunnus alalunga were evaluated by gonad histology in samples of 132 males (58–118 cm fork length, L_F) and 112 females (59–101 cm L_F) that were collected from the western North Pacific Ocean from 2001 to 2006. In the sex ratio examination, males greatly outnumbered females in large adult fish (L_F > 100 cm). Thunnus alalunga exhibited a protracted spawning period from March to September in the waters off eastern Taiwan and the Philippines, and the peak spawning activity occurred in March and April. Minimum sizes associated with the classification of mature fish were 78 and 83 cm L_F for males and females, respectively. In addition, the largest L_F of immature fish were 93 cm for males and 94 cm for females. The spawning frequency estimate in April was 1·7 days. Batch‐fecundity estimates of 21 females (89–99 cm L_F) ranged between 0·17 and 1·66 million eggs (mean ±s.d . = 0·94 ± 0·43). The relative fecundity estimates of the 21 females ranged between 9·2 and 92·4 oocytes g^?1 body mass (mean ±s.d . = 50·5 ± 22·8). The results presented in this study provide increased information regarding this species' reproductive‐related characteristics than are currently available in stock status determinations.     

Batch fecundity and spawning frequency of invasive Hemiculter leucisculus (Basilewsky, 1855) in Erhai Lake,China

Hemiculter leucisculus are multiple spawners with indeterminate fecundity, although previous fecundity estimates have assumed a determinate spawning pattern. Batch fecundity and spawning frequency of H. leucisculus were studied in Erhai Lake, China, in which the fish, as an exotic species, has become a successful colonizer. Spawning frequency was 16.05 %, as determined from the percent of females with postovulatory follicles 12 to 36 h old during the peak reproductive period (from May 27 to August 8). The average interval between spawning was 6.2 days and there were more than 16 total spawning batches during the peak reproductive period. The average batch fecundity (mean ± SD) was 11,934?±?5,921 hydrated oocytes in 40 females (standard length or SL: 9.1–14.4 cm), while relative batch fecundity was 560?±?137 eggs g^?1 wet weight. The total potential annual fecundity was approximately 190,944 oocytes over the whole spawning season and was much higher than the estimated standing stock (31,585 oocytes) of yolked oocytes (36 females, SL: 9.5–16.2 cm) at the beginning of the spawning season in April 2010. This high annual fecundity is likely to have contributed to successful invasion of Erhai Lake by H. leucisculus.     

Sexual cycle and seasonal changes in the ovary of the red mullet, Mullus surmuletus, from the southern coast of Brittany

The reproductive cycle of the red mullet is described on a macroscopic scale in terms of the GSI, HSI and K , and on a microscopic scale in terms of histological changes in the ovary and changes in the oocyte size frequency distribution. On the southern coast of Brittany the red mullet breeds in May and June. During oogenesis, the previtellogenic period lasts 6 months and the secondary phase of vitellogenesis no more than 3 months. When spawning commences the process of vitellogenesis ceases and up to 20% of the vitellogenic oocytes become atretic. Prior to spawning a single batch of oocytes can be seen to be entering secondary vitellogenesis. During the immediate prespawning and spawning periods the existing vitellogenic oocytes mature but there is no recruitment from the stock of previteilogenic oocytes. This results in a gap or hiatus in the oocyte size frequency distribution between previtellogenic and vitellogenic oocytes within which there are very few resting or maturing oocytes. The red mullet appears to be a determinate spawner, in which egg loss through atresia considerably reduces the potential fecundity.     

Reproductive features of the non-native Siganus luridus (Teleostei, Siganidae) during early colonization at Linosa Island (Sicily Strait, Mediterranean Sea)

In July 2003, the finding of a newly settled population of Siganus luridus at Linosa Island (Sicily Strait, Mediterranean Sea) gave us the unusual opportunity to examine the reproductive condition of a Lessepsian migrant during early phases of colonization. Aspects of gonad morphology, fecundity, atresia and oocyte dynamics were investigated by using 43 pioneer specimens collected in concomitance with their first record in the Pelagie Islands. Ovarian development was consistent with the group‐synchronous type, and testicular organization was of the unrestricted spermatogonial testis type, with cystic spermatogenesis. Both males and females had reached final stages of gonad maturation. The rates of follicular atresia were moderate: out of 17 adult females, 10 individuals did not present atretic oocytes; six exhibited <15.1% of secondary growth phase (SGP) oocytes in α‐atresia, while one female presented 45.7% of SGP in α‐atresia. Fecundity estimates did not diverge from what was observed in a reference population along the Lebanese coast. Absolute fecundity ranged from 115 739 to 740 433 oocytes per female (16.5–24.5 cm L_T). Relative fecundity ranged from 1239 to 3162 oocytes g⁻¹, with a mean of 1885 ± 868 oocytes g⁻¹. Our observations indicated that these early settled siganids are reproductively active at Linosa and suggested the forthcoming of self‐maintaining populations across the central Mediterranean area.     

Effect of fish length and nutritional condition on the fecundity of distressed Atlantic cod Gadus morhua from the Baltic Sea

The disappearance of larger individuals and the decrease in individual body condition suffered by Atlantic cod Gadus morhua in the eastern Baltic during the past two decades can be expected to affect the stock reproductive output. To investigate this, female G. morhua were collected during the spawning and pre‐spawning period in 2015?2016. The current individual potential fecundity (F_P) of eastern Baltic G. morhua was estimated and analysed in relation to total length (L_T) and indices of nutritional status such as body condition (K) and hepato‐somatic index (I_H) using generalized linear models. In addition, the current prevalence of atresia and its potential relation to K were investigated. Moreover, a calibration curve to estimate F_P from oocyte diameter, based on the autodiametric oocyte counting method, was established for the first time for eastern Baltic G. morhua and can be used for future fecundity studies on this stock. The results showed that F_P was mainly positively related to fish length, but K and I_H also contributed significantly to the variation in F_P. The model predicted that fish with K = 1·2 have a F_P 51% higher than fish of the same L_T with K = 0·8. The prevalence of fecundity regulation by atresia was 5·8%, but it was found only in fish in the pre‐spawning maturity stage and with low K. Temporal changes in biological features such as the length composition and individual body condition of eastern Baltic G. morhua, should be accounted for when estimating stock reproductive potential.     

Reproductive strategy of the European perch,Perca fluviatilis Linnaeus, 1758 (Osteichthyes: Percidae) in the Anzali wetland,southwest Caspian Sea

Temporal variation in reproductive traits of geographically distributed fish is supposed to take place in response to the spatial and environmental variations. With regard to the wide distribution of the European perch in the northern hemisphere, important reproductive traits such as the initiation and duration of the spawning activity are likely to vary in different latitudinal gradients. In this study, reproductive biology of the European perch, Perca fluviatilis, is described, based on 324 specimens caught in the Anzali wetland (southwest Caspian Sea) between June 2008 and May 2009. The gonadosomatic index, oocyte frequency distribution and histological examination suggested a long vitellogenic process (October to February) and a short spawning season (January and February). The size‐frequency distribution of the oocytes showed that this perch is a species with group‐synchronous ovarian development. Ovarian development occurred only in one clutch of oocytes (700–900 μm oocyte diameter) with no indication of maturation of any subsequent clutch in the spawning season. The average of (realized) fecundity (±SD) was estimated to be 16177 ± 5846 eggs in late vitellogenic stage, which was lower than the potential fecundity (17188 ± 6917 eggs). Histological examination of the gonads revealed the existence of atretic oocytes in early vitellogenic stages (October and November). This investigation highlights the temporal variation in the initiation and duration of the reproductive activity of the European perch in this region compared to other geographical regions. The results emphasize the necessity of specific temporal management in fishing of European perch based on spatial differences in reproductive biology.     

Aspects of the fecundity of the feral catfish,Clarias macrocephalus (Gunther), population obtained from the rice fields used for rice-fish farming,in Malaysia

The catfish, Clarias macrocephalus, harvested from the irrigated rice fields of Malaysia, exhibited spawning activities that are influenced by water level management. The multiple spawning potential of the fish is negated by declining water level and the fish spawned only once per growing season. Although oocyte development started in conjunction with water level increase, subsequent oocyte development did not take place due to low water level. Absolute and relative fecundity also decreased with declining water level, however, mean relative fecundity is relatively constant (17.6–53.7 oocytes g^–1 body weight) and not related to body size. Absolute fecundity ranged between 785 to 14066 oocytes female^–1, whereas relative fecundity is between 7.1 to 83.5 oocytes g^–1 body weight, respectively.     

Reproductive biology of female striped marlin Kajikia audax in the western Pacific Ocean

Length and mass data for 1260 (536 females, 683 males, 41 sex unknown) striped marlin Kajikia audax were collected at the fish markets of Tungkang, Singkang and Nanfangao from July 2004 to September 2010. Of these samples, 534 gonads (236 females and 298 males) ranging from 95 to 206 cm in eye‐to‐fork length (L _EF ) and 8 to 88 kg in round mass (M _R), were collected. Chi‐square tests indicated sex ratios were homogeneous among months in 2004 and 2006–2008, but not in 2005, 2009 and 2010; and there were significant differences in sex ratio by size. The overall sex ratio (R _S ) differed significantly from the expected 0·5. Kajikia audax are sexually dimorphic and the proportions of females increased with size between 140 and 210 cm L _EF . Reproductive activity was assessed using a gonado‐somatic index (I _G ), external appearance of the gonads and histological examination and results indicated that the spawning season occurred from April to August with a peak in June to July. Based on histological observations and the distribution of oocyte diameters, K. audax are multiple spawners and their oocytes develop asynchronously. The estimated length‐at‐50% maturity (L _EF50 ) was c . 181 cm (c . 4·8 years of age) for females. The proportion of reproductively active females in the spawning season with ovaries containing postovulatory follicles (0·27) indicated that they spawned every 3·7 days on average. The hydrated oocyte method estimated mean ± S.D. batch fecundity (F _B ) to be 4·4 ± 2·02 million eggs; average relative fecundity was 53·6 ± 13·9 oocytes g^?1 M _R; and the average annual fecundity was 181·3 ± 48·3 million eggs. The parameters estimated in this study are key information for stock assessments of K. audax in the north‐western and central Pacific and will contribute to the conservation, management and sustainable yield of this species.     

An autofluorescence-based survey of late follicular atresia in ovaries of a teleost fish (Thunnus thynnus)

In this study, the authors examined late atretic follicles in the ovaries of Atlantic bluefin tuna, Thunnus thynnus (Linnæus 1758), at different times of the year using transmitted light and epifluorescence microscopy. Atresia (degeneration and resorption of developing ovarian follicles) is a natural process involved in fecundity downregulation in teleosts and is substantially enhanced in stressful conditions. Early (α and β) atretic stages of yolked oocytes have a relatively short duration in seasonally reproducing species, whereas later (γ and δ) atretic follicles (LAF) persist for longer time in the ovary, serving as a sign of previous vitellogenic activity. LAF can thus be used as reliable markers of maturity during non-reproductive periods. Lipofuscin granules accumulate in the cytoplasm of LAF cells as a result of lysosomal digestion of oocyte components. Taking advantage of the well-known autofluorescent properties of lipofuscins, LAF may be identified in unstained histological sections under fluorescence microscopy using appropriate excitation and emission wavelengths. The authors explore in this study the applicability of fluorescence microscopy to provide a fast and effective method to assess late atresia in fishes. This method may be particularly useful to determine sexual maturity in individuals sampled long after the spawning season, where LAF are difficult to detect in standard histological sections. Furthermore, LAF autofluorescence provides a rapid way to quantify late atresia in fishes using image analysis.     

Cyto-histological examination of post-vitellogenesis and final oocyte maturation in captive-reared striped bass

The ovarian development of captive-reared, striped bass Morone saxatilis was examined during a 10-week period encompassing the spawning season. Vitellogenic oocytes in March had a mean diameter of 838 ± 18 μm and did not grow significantly thereafter. Except from one non-hormone-treated fish, all females failed to undergo final oocyte maturation (FOM) and their ovaries became atretic with the onset of high spring temperatures. A clearing fixative was found useful in identifying early stages of atresia, evident by the absence of the germinal vesicle (GV). Final oocyte maturation of fish treated with gonadotropin-releasing hormone agonist (GnRHa) consisted of two phases. Early FOM lasted from 1 to 3 weeks, and was associated with lipid-droplet coalescence, and displacement of the GV and yolk globules to the peripheral cytoplasm. Late FOM lasted <24h, and consisted of yolk-globule coalescence and GV breakdown (GVBD). Ovulated eggs had completely coalesced lipid and yolk masses, with cortical alveoli lined against the cell wall. Both phases of FOM were associated with significant increases in oocyte diameter. Striped bass oocytes showed important morphological differences compared to oocytes of other members of the Moronidae family, in terms of percentage lipid content, chorion thickness and degree of hydration after ovulation.     

Reproductive biology of female bigeye tuna Thunnus obesus in the western Pacific Ocean

The reproductive biology of female bigeye tuna Thunnus obesus was assessed by examining 888 fish (ranging from 84·9 to 174·4 cm fork length, L_F) caught by Taiwanese offshore longliners in the western Pacific Ocean from November 1997 to November 1998 and November to December 1999 and 258 gonad samples from these fish. The overall sex ratio of the catch during the sampling differed significantly from 0·5, but males were predominant in sizes >140 cm L_F. Reproductive activity (assessed by histology), a gonado‐somatic index, and the size‐frequency distributions of whole oocytes indicated that spawning occurred throughout the year and the major spawning season appeared to be from February to September. The estimated sizes at 50% maturity (L_F50) of females was 102·85 cm (95% c.i .: 90·79–110·21 cm) and the smallest mature female was 99·7 cm L_F. They are multiple spawners and oocytes develop asynchronously. The proportion of mature (0·63) and reproductively active (0·70) females with ovaries containing postovulatory follicles indicated that they spawn almost daily. Batch fecundity for 15 females with the most advanced oocytes (>730 µm) ranged from 0·84 to 8·56 million eggs (mean ± s.d . = 3·06 ± 2·09). The relationships between batch fecundity (F_B, in millions of eggs) and L_F (cm) and round mass (M_R, kg) were (r² = 0·84) and (r² = 0·80), respectively. The parameters estimated in this study are key information for stock assessments of T. obesus in the western Pacific Ocean and will contribute to the conservation and sustainable yield of this species.     

The influence of feeding regime on sexual maturation, fecundity and atresia in first-time spawning turbot

Exposure of adolescent turbot Scophthalmus maximus to low rations during vitellogenesis, covering the 4 months immediately prior to spawning, led to a drop of 70% in mean ovary weight, and was associated with poor growth of the vitellogenic oocytes or, in a third of cases, the absence of vitellogenic oocytes. Exposure to low rations during the recruitment of vitellogenic oocytes, 4–8 months prior to spawning, produced a more variable response. A few of these females produced well-developed vitellogenic oocytes, but in most, vitellogenic oocytes were absent or only poorly developed and in some cases there was a high incidence of atresia. Exposure to intermediate rations throughout oocyte recritment and vitellogenesis also depressed oocyte development. In maturing females the number of non-atretic vitellogenic oocytes (relative potential fecundity) averaged 998 per g of somatic body weight of fish at the start of the spawning season in mid-June. Maturation of the males was not significantly influenced by the dietary regime. The small size of the testes and the low levels of milt production, even in fish fed high rations immediately prior to spawning, showed that reproductive investment in males was much lower than in females.