Comparative study of reproductive biology in single and multiple-spawner cyprinid fish. II. Sex steroid and plasma protein phosphorus concentrations

The dynamics and regulation of oogenesis in single and multiple-spawner cyprinid fish showing group-synchronous oocyte development, was investigated in three species from the River Meuse (Belgium): the roach Rutilus rutilus as a single spawner, and the bleak Alburnus alburnus and the white bream Blicca bjoerkna as multiple spawners. This paper compares the seasonal profiles of sex steroids (oestradiol-17β, testosterone and 17,20β-dihydroxy-4-pregnen-3-one) and plasma alkali-labile protein phosphorus. Different patterns of plasma oestradiol-17β (E2) and plasma protein phosphorus (PPP) have been observed not only between the single and the multiple spawner fish, but also among the two multiple spawners. In roach, two increases of E2 levels were observed. The first occurred in September after a short gonadal quiescent period, and coincided with the increase of the PPP at the onset of exogenous vitellogenesis. The second took place in spring before the spawning season. The low PPP recorded during that period probably reflected its rapid incorporation by the oocytes. In both multiple spawners, highest values of PPP were recorded just before the spawning season. In white bream, the PPP declined progressively once the differentiation of exogenous vitellogenic oocytes was completed before the onset of the spawning season. In bleak, PPP levels remained high throughout the spawning season and corresponded to a sustained oocyte recruitment during the whole of this period. Regardless of the pattern of oocyte growth recruitment, the E2 concentrations were high in both multiple spawner species during the breeding season. In the three species, testosterone levels remained low regardless of the maturation stage (ranging from 0.6 to 13.4ng ml^?1). Except for relatively high concentrations of 17,20βP in roach during final maturation and postspawning stages (20 and 28 ng ml^?1, respectively), low levels of this steroid were measured in these cyprinids, and especially in the multiple spawner fish. The role of this progestogen as the maturation inducing Steroid is discussed.     

Liver Changes Related to Oocyte Growth in Roach,a Single Spawner Fish,and in Bleak and White Bream,Two Multiple Spawner Fish

The objectives of this study were to compare the seasonal hepatic activity and to evaluate whether this activity was related to the dynamics of oocyte recruitment in three cyprinids from the River Meuse (Belgium): the roach Rutilus rutilus as a single spawner, and the bleak Alburnus alburnus and the white bream Blicca bjoerkna as multiple spawners. In roach, hepatosomatic index (HSI) was low during exogenous vitellogenesis. However, the strong development of the rough endoplasmic reticulum (RER), the presence of large mitonchondria with many inner folds, and the depletion of glycogen, lipid droplets and phospholipids indicated intense hepatic activity during that period. Prior to spawning, hepatic activity decreased as indicated by a decrease of the HSI and in the hepatocytes a regression of the RER and the disappearance of the lipid droplets and glycogen. In bleak, the oocyte recruitment occurred continuously throughout the spawning season and the vitellogenic activity of the liver remained intense throughout that period as suggested by high HSI and the ultrastructure of the hepatocytes. In contrast in white bream the vitellogenic activity of the liver decreased during the spawning season as the completion of the vitellogenesis took place prior to spawnings. We concluded that the vitellogenic activity of the liver is strongly related to oogenesis.     

Oogenesis in the bluefin tuna,Thunnus thynnus L.: a histological and histochemical study

Histology and histochemistry are useful tools to study reproductive mechanisms in fish and they have been applied in this study. In the bluefin tuna, Thunnus thymus L., oocyte development can be divided into 4 principal phases based on the morphological features of developing oocytes and follicles. The primary growth phase includes oogonia and basophilic or previtellogenic oocytes classified as chromatin-nucleolus and perinucleolus stages. The secondary growth phase is represented by vitellogenic oocytes at early (lipid globule and yolk granule 1), mid (yolk granule 2) and late (yolk granule 3) vitellogenesis stages. The maturation phase involves postvitellogenic oocytes undergoing maturation process. During the spawning period, both postovulatory follicles, which indicate spawning, and atretic follicles can be distinguished in the ovary. Carbohydrates, lipids, proteins and specially those rich in tyrosine, tryptophan, cystine, arginine, lysine and cysteine, as well phospholipids and/or glycolipids and neutral glycoproteins were detected in yolk granules. Moreover, affinity for different lectins (ConA, WGA, DBA and UEA) was detected in vitellogenic oocytes (yolk granules, cortical alveoli, follicular layer and zona radiata), indicating the presence of glycoconjugates with different sugar residues (Mannose- Man- and/or Glucose -Glc-; N-acetyl-D-glucosamine- GlcNAc- and/or sialic acid- NANA-; N-acetyl-D-galactosamine- GalNAc-; L-Fucose -Fuc-). Histochemical techniques also demonstrated the presence of neutral lipids in globules (vacuoles in paraffin sections) and neutral and carboxylated mucosubstances in cortical alveoli. By using anti-vitellogenin (VTG) serum, immunohistochemical positive results were demonstrated in yolk granules, granular cytoplasm and follicular cells of vitellogenic oocytes. Calcium was also detected in yolk granules and weakly in follicular envelope. In females, the gonadosomatic index (GSI) increased progressively from May, during early vitellogenesis, until June during mid and late vitellogenesis, where the highest values were reached. Subsequently, throughout the maturation-spawning phases (July), GSI decreased progressively reaching the minimal values during recovering-resting period (October).     

Reproductive biology ofAwaous guamensis, an amphidromous Hawaiian goby

Synopsis Spawning season, size at first reproduction, oocyte maturation, and fecundity ofAwaous guamensis, an amphidromous Hawaiian goby, were studied from June 1989 through May 1991 in the Wainiha River, Kau'ai, Hawai'i. Female fish larger than 73 mm standard length (SL) had mature gonads from August through December in 1989 and 1990. Gonadosomatic index (GSI) values for mature females ranged from 0.2 to 14.5 during the spawning season. Male fish larger than 64 mm SL had elevated GSI values from June 1989 through December 1989 and from August 1990 through December 1990. Mature sperm were found in two male fish collected in January and February. GSI values for mature males ranged from less than 0.01 to 4.0 in the spawning season. Size-frequency distributions of measurements of vitellogenic oocyte diameters and microscopic observations of oocytes indicated this species has group-synchronous oocyte development. Ovarian maturation stages examined over a 29-month period suggest that members of the stock spawned at different times within the spawning season, although mass spawning events have been documented for this species. Estimates of clutch sizes from nests measured in situ were comparable to estimates of potential fecundity from in vitro examination of ovaries, and indicated that female fish deposited an entire clutch during a spawning event. No evidence for multiple spawning by an individual fish in a single season was found. However, microscopic observations of brown bodies in some ovaries suggested that individual fish probably spawn more than once in a lifetime.     

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.     

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.     

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.     

Gross and histological observations of ovarian development and related condition changes in American plaice

The largest increase in gonadosomatic index of American plaice Hippoglossoides platessoides occurred between January and February. Hepatosomatic index was lowest in the spawning season, suggesting cessation of feeding activity during spawning, but rose immediately in July when condition factor recovered from its low spawning level. Increases in white muscle water content during the spawning season suggests protein depletion. Conservation of some white muscle fibres is suspected, but no regular pattern in area conserved was found. Gonad development was advanced in December and in April some gonads were showing evidence of spawning. Ovaries from spawning females showed hydration patterns and histological detail consistent with batch or serial spawning strategy. Evidence of recent spawning activity, including the presence of postovulatory follicles, was found in ovaries that also contained oocytes undergoing exogenous vitellogenesis. Oocyte size frequency distributions show a gap in size between cortical alveoli and vitellogenic oocytes prior to spawning which does not exist during the spawning season. This may represent the ability of plaice to push oocytes through vitellogenesis from a previtellogenic condition during the spawning period. The increased white muscle moisture during June and July may reflect the use of muscle protein to bring oocytes through vitellogenesis for the current spawning season.     

The dynamics of oocyte growth during vitellogenesis in the rainbow trout (Oncorhynchus mykiss)

This report describes the dynamics of oocyte growth during vitellogenesis in a population of virgin female rainbow trout. Indices of ovarian development increased dramatically during the period of study: the gonadosomatic index (GSI) increased over 50-fold, reaching a peak of 20 just before ovulation; the mean oocyte diameter increased from less than 1 mm to 5.4 mm; and plasma levels of vitellogenin increased from less than 1.5 mg/ml to 25 mg/ml. There were no changes in the numbers of developing oocytes (measuring 0.5 mm or greater in diameter) from the time when the majority of oocytes undergoing secondary development had entered vitellogenesis in August to ovulation in February (averaging 4000 oocytes per fish). The increase in ovary weight during vitellogenesis was, therefore, due to an increase in the size of oocytes rather than to recruitment of more maturing oocytes. The numbers of vitellogenic oocytes in the ovary during the entire study also suggested that atresia of vitellogenic oocytes does not play a prominent role in determining fecundity. During early vitellogenesis, the volume of maturing oocytes within an ovary varied by as much as 250-fold. From September onwards, when all oocytes to be ovulated that season had entered vitellogenesis, a gradual uniformity in size began to develop, such that at ovulation, in February, all the eggs were very similar in size (there was less than a 2-fold variation in volume). The pattern of growth of oocytes in an ovary during vitellogenesis suggests that growth between oocytes is closely coordinated.     

Comparative study of sexual cycles, oogenesis and spawning of two Soleidae, Solea lascaris (Risso, 1810) and Solea impar (Bennet, 1831), on the western coast of Brittany

Two sand sole species, Solea lascaris and Solea impar , are common on the western coast of Brittany, France. Their reproduction (oogenesis and spawning) has been studied by following the gonadosomatic index, histological changes in ovary development, and oocyte diameter distribution. Both species have a prolonged spawning season, from spring to late summer. Each mature female breeds twice during the spawning season: in May and July for 5. lascaris , and in early June and July for 5. impar. Throughout the spawning season, several oocyte batches undergo vitellogenesis and there is a continuous recruitment of immature or primary oocytes into vitellogenesis. The two species are thus serial synchronous spawners. This, together with oocyte atresia late in the season (especially noticeable in S. impar ) makes fecundity estimation difficult.     

Reproductive cycles in a reared strain of the mummichog, a daily spawner

Annual, lunar, and diel samplings were taken from a strain of mummichog (Arasaki strain) reared in outdoor tanks under natural conditions, to examine gonadal maturity. Gonads of yearling fish were quite immature in September. During late autumn and winter, a gradual increase in the GSI of both sexes was observed, and the growth of cortical alveolus phase oocytes in females and basal spermatogenesis in males progressed. In late February, a rapid increase in the GSI of both sexes, vitellogenesis in females, and active spermatogenesis in males, occurred. The spawning period of the yearling fish was from late March to August judging from the presence of milt-producing males and ovulated females. The spawning period of the underyearling fish started in the same month as the yearlings, but terminated 1 month earlier. Plasma levels of oestradiol-17 β (E₂) in females and testosterone in males were high during the spawning period in the yearlings. In the underyearlings, however, E₂ levels peaked in early spring, and declined in the latter part of the spawning period. Neither a lunar nor semilunar cycle was evident in the reproductive activity of this fish, which proved to be a typical daily spawner. Females showed an apparent daily reproductive cycle; oocyte maturation commenced at about 1200 hours, germinal vesicle breakdown (GVBD) occurred at about 2400 hours, and ovulation was completed by 2400 hours, 24 h after GVBD. Such clear annual and daily reproductive cycles make this strain of mummichog a suitable model for the study of environmental and endocrine regulation of reproductive cycles in marine and estuarine teleosts.     

Reproductive biology of the armoured catfish Loricariichthys castaneus (Castelnau, 1855) in Lajes reservoir,southeastern Brazil

Sex ratio, gonadal development, breeding season and fecundity of the armoured catfish Loricariichthys castaneus were described to assess its reproductive strategy in a Brazilian tropical reservoir. In total, 226 specimens (199 females and 27 males) were captured from September 2005 to August 2006 and examined in the laboratory. Females outnumbered males and achieved sizes larger than 330 mm TL. Oocyte development, determined by histological analysis, was asynchronous with oocyte size, ranging from pre‐spawning (27–270 μm) to spawning (243–3460 μm), followed by a sharp decrease in the mean oocyte diameter postspawning (590–730 μm) as the spawning proceeded. Spawning occurred throughout most of the year, peaking in August–September and reaching a low in April–May, according to variations in GSI and frequencies of stages of gonadal development. Batch fecundity ranged from 242 to 833 vitellogenic oocytes (relative fecundity = 2.27 oocytes g^?1), averaging 483 oocytes, and was positively related to gonad weight (P = 0.00003). Oocyte diameters ranged from 0.027 to 5.59 mm, with vitellogenic diameters ranging from 2.08 to 5.59 mm. Continuous development of oocytes throughout the year suggests that L. castaneus presents indeterminate fecundity and is a batch‐spawner. These attributes, associated with parental care and a wide reproductive period, correspond to an equilibrium strategy that has proved to be effective in the Lajes reservoir.     

The effects of thermal effluent exposure on the gametogenesis of female fish

High temperature in Swedish and Lithuanian thermal effluent areas influenced gametogenesis of female perch Perca fluviatilis , roach Rutilus rutilus and pike Esox lucius negatively, indicating reduced reproductive capacity. Oocyte atresia started during vitellogenesis in autumn, and was often followed by asynchronous egg cell development. Among other anomalies, multi-nucleus oocytes and hermaphroditism were observed. No significant impact was seen in silver bream Blicca bjoerkna . Ruffe Gymnocephalus cernuus reacted by a tendency to produce an additional mature oocyte generation during the spawning period. Ovaries in roach from coastal areas were often infected by a microsporidian parasite Pleistophora mirandellae , causing severe damage to the gonad. Parasites were also detected in pike, but neither in perch nor in lake populations of roach. Fish living in open coastal environments did not avoid impact by moving out of the heated areas. There seems to be a conflict in some temperate fish between temperature preference behaviour and safeguarding normal reproduction.     

Histological and histochemical study of female germ cell development in the dusky grouper Epinephelus marginatus (Lowe, 1834)

The developmental stages of female germ cells were analysed in a wild population of the protogynous teleost Epinephelus marginatus (Lowe, 1834). 321 wild dusky grouper females were collected in the South Mediterranean Sea during the spawning season and their ovaries analysed using histological and histochemical techniques. Oocyte morphology, nucleus-cytoplasm ratio (N/C) range, location and movements of cytoplasmic inclusions during primary growth, vitellogenesis and final oocyte maturation were described. The distribution of proteins, lipids and carbohydrates through oocyte development was also investigated in 50 females. Lipid vesicles appeared firstly in the mid ooplasm of oocytes larger than 130 microm, at the beginning of the secondary growth phase. Immediately afterwards, small carbohydrate granules (PAS and Alcian blue positive) appeared before the occurrence of the first yolk granules. Tyrosine-enriched proteins were especially evidenced in the zona radiata interna of late vitellogenic oocytes. Specific lectin binding patterns reflected characteristic differences in the content and distribution of specific sugar moieties expressed in the oocytes during vitellogenesis and final maturation. At the end of vitellogenesis and during final maturation, follicular cells, zona radiata, and cortical alveoli were characterised by a strong increase of specific binding for WGA.     

Occurrence of cyprinids in fish ladders in relation to flow

To analyse the relationship between spawning migrations and flow, cyprinids from ten fish ladders of the Elbe River (Czech Republic) were collected during the period of spawning migrations (April–June) in 1996, 2000 and 2001. Number of species and individuals in the fish ladders increased with increasing flow up to medium values, followed by further decrease during high flows, whilst length of individuals increased with increasing flow. Moreover, occurrence of four the most numerous species (bleak, roach, silver bream and barbel) showed species specific differences in response to flow variability. In early spring, the highest number of bleak in fish ladders occurred during the low flow, whilst later increased with increasing flow. Occurrence of roach and silver bream achieved the maximum during the medium flow values across the whole studied period. In opposite, occurrence of barbel increased with increasing flows during the whole period. Furthermore, it can be assumed that occurrence of fish in ladders was associated not only with spawning migrations but also reflected shifts in local fish distribution in response to flow variability in the main channel.     

The multiple spawning pattern of weakfish in the Chesapeake Bay and Middle Atlantic Bight

Weakfish Cynoscion regalis were collected from commercial fisheries in the Chesapeake Bay and the Middle Atlantic Bight (n=4380) during 1989–1992 and their reproductive biology assessed using the gonadosomatic index, macroscopic gonad stages, oocyte diameter distributions, microscopic whole oocyte analysis and histology. Sex ratios were approximately 3:1, females to males, in 1990–1992. Most fish (90%) attained sexual maturity by age 1 and at a small size. Estimated mean length at first maturity was: 164mm total length (TL) for males, and 170 mm TL for females. Weakfish spawn within the Chesapeake Bay, as far north as the Virginia/ Maryland border. Although spawning occurred during May–August and gonad development and initiation of spawning was synchronous, cessation of spawning was asynchronous. There was no indication that older fish exhibited a more extended spawning season than younger fish. Weakfish are multiple spawners with indeterminate fecundity. Oocyte development is asynchronous with oocytes of all stages being present in developed ovaries. Because of the complex and dynamic weakfish ovarian cycle, typical methods of assessing reproduction, such as the GSI and macroscopic gonad stages, are inadequate for this species if not used in conjunction with more detailed methods such as histology.     

Spawning frequency, fecundity, egg weight and spawning type of silver pomfret, Pampus argenteus (Euphrasen) (Stromateidae), in Kuwait waters

Silver pomfret, Pampus argenteus, were collected by fishing with drift gillnets on one spawning ground in Kuwait waters during 1998–2000. Fish size frequency, sex ratio, maturation cycle, spawning frequency, fecundity and egg weight were assessed. The length–weight relationship differed between sexes whereby females were significantly bigger than males. Spawning started in mid‐May and continued until early October. During this time the water temperature ranged from 26.0 to 32.8°C, salinity was ? 39.0‰ and water depth ranged between 5 and 12 m. Large females spawned earlier than young spawners and the overall percentage of males during the spawning period was 70.3%. Spawning occurred after 13.00 h, with peak spawning between 15.00 and 18.00 hours during outgoing tide. Mean daily spawning frequency amounted to 63.2%. Spawning activity was found to be associated with the lunar cycle and spawnings were concentrated during the first and third quarters of the moon period, indicating a semilunar reproduction cycle. It was concluded that a female would spawn at least six times during the season. No change was observed in relative fecundity during the peak spawning season (June–August). Average relative batch fecundity was 176.3 eggs g^?1 somatic weight (SW), corresponding to a relative total fecundity of 1058 eggs g^?1 SW, which is 1.5 times higher than estimates obtained from counting the standing stock of oocytes. Bigger fish produced heavier eggs and the egg weight decreased as the spawning season progressed. Based on gonadal cycles, oocyte size frequency distribution and total fecundity, we concluded that silver pomfret is a multiple batch spawner with indeterminate fecundity.     

Gonadal development and sex steroids in a female Arctic charr broodstock

Gonadal development and plasma levels of sex steroids were investigated in female Arctic charr at 3-week intervals over a 12-month period. Circulating levels of oestradiol-17β (E₂), testosterone (T) and 17,20β-dihydroxy-4-pregnen-3-one (17,20β-P) were measured by radioimmunoassay, and gonadal status assessed through histological examination and measurement of gonadosomatic index (GSI) and frequency distribution of oocyte size-classes. Gonadal recrudescence during March-July was characterized by modest but insignificant increases in plasma levels of E₂ (2–4 ng ml^?1) and T (2–5 ng ml^?1) and recruitment of oocytes into yolk accumulation. Only a small and insignificant rise in GSI and no apparent increase in oocyte diameter occurred during this period, indicating that the rate of yolk formation and oocyte growth was low. Following transformation from stage V (peripheral yolk granule stage) to stage VI (yolk granule migration stage) in late July, the vitellogenic oocytes entered a phase of rapid growth which resulted in a marked rise in GSI until ovulation commenced in late September. Gonadal growth during this period was accompanied by increases in plasma levels of E₂ and T which peaked at 11 ± 1 (mid-August) and 71 ± 5ng ml^?1 (late September), respectively. The levels of both steroids dropped rapidly during final maturation and ovulation, followed by a surge in plasma levels of 17,20β-P which peaked at an average of 74 ± 17 ng ml^?1 in early October. All three steroids returned to basal levels within a month after ovulation, and all steroids, except E₂, remained low until March of the following year. A slight increase in E₂ detected in February and March during the second season may have been associated with recruitment into vitellogenesis of a new generation of oocytes. It is suggested that the abrupt increase in vitellogenesis in late July may reflect a condition-dependent decision to proceed with maturation, once the energy reserves have been repleted during spring-early summer.     

Oogenesis and plasma levels of sex steroids in cultured females of brown trout (Salmo trutta linnaeus, 1758) in Chile

Naturalized brown trout populations in Chile are a valuable genetic resource with aquaculture potential. The oogenesis of a three-year-old brown trout cultured population was studied in southern Chile. Gonadosomatic index (GSI), oocyte growth, gonadal microscopic characteristics, and plasma levels of estradiol-17beta (E2), testosterone (T), and 17alpha-hydroxyprogesterone (17alpha-HP) were measured bimonthly for a nine-month period before spawning. The maximum GSI level (22%) was similar to that described for other salmonids, although it was reached in May, more than one month before the population started spawning. Oocyte growth increases strongly from January when diameter reaches more than 1 mm. The vitellogenic period (six-seven months) is consistent with the long vitellogenesis, described for salmonid females maturing at three years old. E2 shows a slow increase from November, reaching its peak value in March (65.2+/-0.7 ng/ml), during maximal vitellogenic activity. T increases as oogenesis progresses, reaching a maximum of 90+/-20 ng/ml during May, and falling considerably during ovulation. Following a typical pattern of progestogens in salmonid oogenesis, 17alpha-HP stays at basal levels during most of oogenesis, but experiences a strong surge (2.0+/-0.4 ng/ml) just before ovulation.