Sex change in the protandrous black porgy, Acanthopagrus schlegeli: a review in gonadal development, estradiol, estrogen receptor, aromatase activity and gonadotropin.

Black porgy, Acanthopagrus schlegeli Bleeker, a marine protandrous hermaphrodite, is functional male for the first two years of life but begins to sexually change to female after the third year. Testicular tissue and ovarian tissue was separated by connective tissue in the bisexual gonad. This sex pattern provides a very good model to study the endocrine mechanism of sex change in fish. The annual profiles of plasma estradiol, vitellogenin and 11-ketotestosterone concentrations in males were significantly different from those in the three-year-old females. Significantly high levels of plasma estradiol during the prespawning/spawning season and low levels of plasma 11-ketotestosterone during the spawning season were observed in the inversing females. No difference of plasma testosterone levels was observed in males and females. Oral administration of estradiol stimulated high levels of gonadal aromatase activity, plasma gonadotropin II levels and sex change in the two-year-old fish. Exogenous estradiol administered for 5-6 months induced a reversible sex change in one- and two-year-old fish. The sensitive period for estradiol treatment of sex change is from early prespawning to spawning season. Implantation with testosterone for more than a year could not block the natural sex change in three-year-old fish. Exogenous aromatase inhibitors (1,4,6-androstatriene-3,17-dione or fadrozole) suppressed aromatase activity in the brain. Oral administration with aromatase inhibitors for a year further inhibited the natural sex change in three-year-old black porgy and all fish became functional male with spermiation. Estrogen receptor alpha gene in the ovarian tissue of bisexual gonad is significantly less expressed than that in the vitellogenic ovary of female on the basis of reverse-transcription polymerase-chain reaction. There was no difference in the annual profiles of the plasma gonadotropin II levels in the males and natural inversing females. Plasma gonadotropin II levels were significantly higher in estradiol-treated group than those in the control. It is concluded that estradiol, aromatase activity and estrogen receptor in the ovarian tissue play an important role in the natural and controlled sex change in black porgy. The association of gonadotropin and sex change in black porgy is not clear.

Europa's differentiated internal structure: inferences from four Galileo encounters.

Radio Doppler data from four encounters of the Galileo spacecraft with the jovian moon Europa have been used to refine models of Europa's interior. Europa is most likely differentiated into a metallic core surrounded by a rock mantle and a water ice-liquid outer shell, but the data cannot eliminate the possibility of a uniform mixture of dense silicate and metal beneath the water ice-liquid shell. The size of a metallic core is uncertain because of its unknown composition, but it could be as large as about 50 percent of Europa's radius. The thickness of Europa's outer shell of water ice-liquid must lie in the range of about 80 to 170 kilometers.

Distribution of rock, metals, and ices in Callisto.

Radio Doppler data from a single encounter (C3) of the Galileo spacecraft with Callisto, the outermost Galilean moon of Jupiter, indicated that Callisto was probably undifferentiated. Now, similar data from a second encounter (C9) corroborate this conclusion, but more accurate data from a third encounter (C10) indicate that the rock and ice within Callisto have partially, but not completely, separated. Callisto may be differentiated into a rock-metal core less than 25 percent of Callisto's radius, an outer layer of clean ice less than 350 km thick, and a middle layer of mixed rock and ice. Models in which ice and rock are mixed all the way to the center of Callisto are also consistent with the data.

Europa's differentiated internal structure: inferences from two Galileo encounters.

Doppler data generated with the Galileo spacecraft's radio carrier wave during two Europa encounters on 19 December 1996 (E4) and 20 February 1997 (E6) were used to measure Europa's external gravitational field. The measurements indicate that Europa has a predominantly water ice-liquid outer shell about 100 to 200 kilometers thick and a deep interior with a density in excess of about 4000 kilograms per cubic meter. The deep interior could be a mixture of metal and rock or it could consist of a metal core with a radius about 40 percent of Europa's radius surrounded by a rock mantle with a density of 3000 to 3500 kilograms per cubic meter. The metallic core is favored if Europa has a magnetic field.

Gravitational evidence for an undifferentiated Callisto.

Before the arrival of the Galileo spacecraft at Jupiter, models for the interior structure of the four galilean satellites--Io, Europa, Ganymede and Callisto-ranged from uniform mixtures of rock and ice (that is, undifferentiated objects) or rocky cores surrounded by a mantle of water ice. Now it appears that Io has a large metallic core and that Ganymede is strongly differentiated, most probably into a three-layer structure consisting of a metallic core, a silicate mantle and a deep outer layer of ice. Direct information on the interior structure of Callisto determined from previous spacecraft fly-bys was essentially limited to an estimate of the mean density being intermediate between pure ice and pure rock. Here we report measurements of Callisto's gravitational field which reveal that, in contrast to Io and Ganymede, this galilean satellite is most probably a homogeneous object consisting of a solar mixture of 40% compressed ice and 60% rock (including iron and iron sulphide). Callisto's undifferentiated state is consistent with the apparent lack of an intrinsic magnetic field, and indicates that the outermost galilean satellite has not experienced a heating phase sufficiently high to separate its rock and metal components from the lighter ices.

The endocrine mechanism of sex reversal in the protandrous black porgy, Acanthopagrus schlegeli: a review.

Black porgy, Acanthopagrus schlegeli Bleeker, a marine protandrous hermaphrodite, is a functional male for the first 2 years of life but begin to sexually reverse to female after the third year. This sex pattern provides a very good model to study the mechanism of sex reversal in fish. The gonad at 5 month of age consisted of testicular tissue with few primary oocytes at 5 month of age. The ovarian tissue became dominant at 18 months of age during the non-spawning season. Testicular and ovarian tissues were separated by connective tissue. Plasma estradiol-17 beta(E2), vitellogenin and 11-ketotestosterone (11-KT) profiles in males were significantly different from those in the 3-year-old reversing females. Peak levels of plasma E2 in the reversing females occurred during the early prespawning season (in October). Lower levels of plasma E2 were, however, observed in the males. Plasma 11-KT levels significant decreased but no changes of plasma testosterone were detected in the reversing females. Exogenous E2 suppressed the testicular development but induced the gonadal aromatase activity, ovarian development and sex reversal in 2-year-old black porgy. Exogenous T and LHRH analog did not have effects on the sex reversal. Higher concentrations of pituitary GtH II and mRNA of GtH II-beta subunit were detected in the reversed females. These data suggested that E2 and gonadal aromatase closely associated to the occurrence of sex reversal. A working model of the sex reversal in black porgy is proposed.

Characterization of vitellogenin induced by estradiol-17ß in protandrous black porgy,Acanthopagrus schlegeli.

The objectives of this study were to isolate and characterize vitellogenin from plasma of estradiol-treated protandrous black porgy,Acanthopagrus schlegeli. Vitellogenin concentrations in plasma measured by a validated enzyme-linked immunosorbent assay (ELISA) were also compared. Two-year-old black porgy (n=20) were fed with estradiol-17ß (4 mg kg(-1) of feed). Plasma was collected for purification of vitellogenin. Two forms of vitellogenin were found in plasma after chromatography on Sepharose CL-2B column and hydroxylapatite, and polyacrylamide gel electrophoresis. Black porgy vitellogenins are phospho-lipo-glycoproteins based on their chemical staining properties. The apparent molecular weights of the two forms of vitellogenin were 636 kDa and 321 kDa, respectively. The amino acid composition of purified vitellogenin was also analyzed after acid hydrolysis. The presence of immunoreactive vitellogenin was identified in the plasma and mucus extract from control and estradiol-induced females on the basis of Western blotting. Serial dilution of the plasma and mucus extract taken from estradiol-induced black porgy showed reactivity to an antiserum against lipovitellin in the ELISA, whereas mucus extract and plasma from male fish did not. Significantly higher concentrations of plasma vitellogenin were detected in estradiol-stimulated black porgy than in the control males.

Stimulation of spermatogenesis or of sex reversal according to the dose of exogenous estradiol-17 beta in juvenile males of protandrous black porgy, Acanthopagrus schlegeli.

The objective was to investigate the effects of various doses of estradiol-17 beta (E2) on gonadal development, gametogenesis, and plasma levels of sex steroids in 1-year-old protandrous male black porgy, Acanthopagrus schlegeli. Juvenile black porgy were divided into four groups, fed with control diet or diet mixed with E2 (0.25, 1.0, or 4.0 mg/kg feed) for 7 months, respectively. Significantly higher gonadosomatic index was observed in the 0.25- and 1.0-mg E2 groups compared to that in the control and 4.0-mg E2 groups during the spawning season. E2 at 4.0 mg completely suppressed spermiation, while lower doses of E2 (0.25 and 1.0 mg) delayed spermatogenesis but increased the number of spermiating fish and spermiation volume. Sperm concentrations were similar in spermiating black porgy. At 10 months and older, gonads in all the groups except E2 (4.0-mg)-treated fish underwent an intensive spermatogenesis while spermatids were not observed until 11 and 12 months in the 0.25- and 1.0-mg E2 groups, respectively. Primary oocytes appeared in the 10- and 11-month-old gonads in testicular tissue of the 1.0- and 4.0-mg E2 groups. After 5 months of 4-mg E2 treatment, testicular tissue regressed and ovarian tissue with primary oocytes developed. Elevated levels of plasma E2 were observed only in the 4.0-mg E2 group while low levels of E2 and testosterone (T) were observed in the control and other E2-treated groups. Significantly higher levels of plasma 11-ketotestosterone (11-KT) were observed in the 0.25- and 1.0-mg E2-treated groups during the spawning season. Plasma vitellogenin levels showed a dose-dependent effect of E2 treatment. The present data suggest that exogenous E2 could stimulate the development of either testicular or ovarian tissues, depending on the dosage of E2. Plasma 11-KT but not T concentrations closely correlate with testicular development and spermiation in black porgy.

Estradiol-17ß suppresses testicular development and stimulates sex reversal in protandrous black porgy, Acanthopagrus schlegeli.

Two year old black porgy (Acanthopagrus schlegeli) fed a diet containing 4.0 mg kg(-1) of estradiol-17ß (E2) for 5 months had significantly lower GSI than the control group during the spawning season. E2 suppressed testicular development, spermiation and plasma testosterone (T) and 11-ketotestosterone (11-KT) and stimulated ovarian development, vitellogenesis and sex reversal. Spermiation in the control group occurred in January and February with the concentrations of 1.08-1.36 × 10(10) sperm ml(-1) of milt. Higher plasma T and 11-KT, but lower E2 levels were detected in the spermiating fish (control group). Higher plasma E2 levels were detected in the sex reversing black porgy during the pre-spawning season. A sharp rise in plasma 11-KT and a drop in T levels were detected in spermiating fish (control group) from January to February. Plasma 11-KT levels correlated with the testicular development and spermiation. The data suggest that E2 plays an important role in controlling the sex reversal of black porgy.