Survey of reproductive physiology on Epinephelus coioides in Khozestan province, Persian Gulf

Epinephelus coioides (family serranidae) is protogynous. This species is one of the most important fishes in food chain of marine proteins of persian Gulf. Therefore knowing about the reproductive biology and physiology of this species is an important role on aquaculture procedures. Monthly samples of Epinephelus coioides were obtained in khozestan Bahrekan province from 2001 to 2002 for annual variation of base line of reproductive hormone. The hormones such as: 17-B estradiol, Testosteron, Progesterone, Gonadotropin I ,II GTHI, II) and cortisol have assayed and also different stages of gonads from the histological point of view were studied by light and electron microscope. Aditional to morphometric and fecundity measurements, the important factors such as : Gonadosomatic index (GSI) Hepatosomotic index (HSI) and Condition factor (KF) were also studied. Environmental factors such as temperature, salinity, photoperiod and pH were analyzed for the determination of effective factors responsible for the changes of reproductive cycles. The flactmation of estroid hormones and gonadotropines show a significant variation in different stages of maturation, e.g 17-B estradiol's concentration in the third stages, GTH II in fourth stages of sexual maturation or final oocyte maturation, plasma Testosteron in post ovulation and Progesterone during maturation indicates the highest levels of above mentioned hormones. The total calcium concentration was high in all year. calcium concentration was correlated with GTH II synthesis and increases with GTH II in June. 17-B estradiol concentration was also correlated with GSI. The high concentration of cortisol throughout the year was an index of stress and development of ovary maturational processes. This species was protogynous synchronous hermaphrodites , and belongs to annual spawning species, being monandric. The sexual transition was found to occure in individuals of 51.2- 105 cm in length. GSI and HSI level confirms the time of spawning period is in April- June. Electrone microscopic studies of gonad tissues showed some changes in mitochondria and endoplasmic reticulum in the post ovulation, maturation and post spawning periods. During the monthly sampling the biochemistry of tissues variations indicated decrease in protein and lipid content, but an increase in water content of spawning fishes which was correlated to the maturation of Epinephelus coioides . sex ratio indicative of higher frequences of females to males during monthly sampling periods. The females were smaller than males in sizes, therefore the females lived in 8-15m depth, but males were living in upper limits of depth. The results indicated that the temperature was the most effective parameter in reproductive cycle of Epinephelus coioides and the mean 24°c was a convenient temperature for spawning. Photoperiod was the second effective. factor on the reproductive cycle for this species. It seemed that the increase in the photoperiod between January to May caused a development of the oocyte. Regarding to the results of this research, it seems that the period of spawning in Epinephelus coioides is in May- June and the aquaculture procedure of Epinephelus coioides could be performed in the above mentioned periods.

Maturation of Nineteen Species of Finfish off the Northeast Coast of the United States, 1985-1990

Observations on maturation stages of nineteen species of economically important finfish off the Northeast coast of the USA were analyzed to examine relationships between fish size or age, and maturity. Maturation schedules and median lengths (L50) and ages (A50) at maturation were derived by fitting the logistic model to the observed proportions. Analyses were generally restricted to observations from 1985 to 1990 obtained during stratified random bottom trawl surveys conducted in spring and autumn by the Northeast Fisheries Science Center and the Commonwealth of Massachusetts Division of Marine Fisheries in waters of the continental shelf from Nova Scotia to Cape Hatteras, North Carolina. Butterfish, Peprilus triacanthus, attained sexual maturity at the smallest median length (11.4 cm, males) and pollock, Pollachius virens, at the highest (41.8 em, males). Median length at maturity for gadiforms ranged from 22.2 to 41.8 em. Within the pleuronectiforms, median length at maturity ranged from 19.1 to 30.4 cm. Median lengths for the pelagic and miscellaneous demersal species were in the same ranges as the pleuronectiforms. Butterfish also attained sexual maturity at the youngest median age (0.9 yr, both sexes) whereas redfish, Sebastes fasciatus, were the latest to mature (5.5 yr, both sexes). For gadids, the median age at maturity ranged from 1.3 to 2.3 yr. Within the pleuronectiforms, median age at maturity ranged from 1.3 to 4.4 yr and, for pelagic species, from 0.9 to 3.0 yr. Median lengths and ages for many species are lower than those reported in earlier studies of the same general region of the Northwest Atlantic. (PDF file contains 72 pages.)

Estimates of body sizes at maturation and at sex change, and the spawning seasonality and sex ratio of the endemic Hawaiian grouper (Hyporthodus quernus, F. Epinephelidae)

A case study of the reproductive biology of the endemic Hawaiian grouper or hapu’upu’u (Hyporthodus quernus) is presented as a model for comprehensive future studies of economically important epinephelid groupers. Specimens were collected throughout multiple years (1978–81, 1992–93, and 2005–08) from most reefs and banks of the Northwestern Hawaiian Islands. The absence of small males, presence of atretic oocytes and brown bodies in testes of mature males, and both developed ovarian and testicular tissues in the gonads of five transitional fish provided evidence of protogynous hermaphroditism. No small mature males were collected, indicating that Hawaiian grouper are monandrous (all males are sex-changed females). Complementary microscopic criteria also were used to assign reproductive stage and estimate median body sizes (L50) at female sexual maturity and at adult sex change from female to male. The L50 at maturation and at sex change was 580 ±8 (95% confidence interval [CI]) mm total length (TL) and 895 ±20 mm TL, respectively. The adult sex ratio was strongly female biased (6:1). Spawning seasonality was described by using gonadosomatic indices. Females began ripening in the fall and remained ripe through April. A February–June main spawning period that followed peak ripening was deduced from the proportion of females whose ovaries contained hydrated oocytes, postovulatory follicles, or both. Testes weights were not affected by season; average testes weight was only about 0.2% of body weight—an order of magnitude smaller than that for ovaries that peaked at 1–3% of body weight. The species’ reproductive life history is discussed in relation to its management.

Relative pleopod length as an indicator of size at sexual maturity in slipper (Scyllarides squammosus) and spiny Hawaiian (Panulirus marginatus) lobsters

Body size at gonadal maturity is described for females of the slipper lobster (Scyllarides squammosus) (Scyllaridae) and the endemic Hawaiian spiny lobster (Panulirus marginatus) (Palinuridae) based on microscopic examination of histological preparations of ovaries. These data are used to validate several morphological metrics (relative exopodite length, ovigerous condition) of functional sexual maturity. Relative exopodite length (“pleopod length”) produced consistent estimates of size at maturity when evaluated with a newly derived statistical application for estimating size at the morphometric maturation point (MMP) for the population, identified as the midpoint of a sigmoid function spanning the estimated boundaries of overlap between the largest immature and smallest adult animals. Estimates of the MMP were related to matched (same-year) characterizations of sexual maturity based on ovigerous condition — a more conventional measure of functional maturity previously used to characterize maturity for the two lobster species. Both measures of functional maturity were similar for the respective species and were within 5% and 2% of one another for slipper and spiny lobster, respectively. The precision observed for two shipboard collection series of pleopod-length data indicated that the method is reliable and not dependent on specialized expertise. Precision of maturity estimates for S. squammosus with the pleopod-length metric was similar to that for P. marginatus with any of the other measures (including conventional evidence of ovigerous condition) and greatly exceeded the precision of estimates for S. squammosus based on ovigerous condition alone. The two measures of functional maturity averaged within 8% of the estimated size at gonadal maturity for the respective species. Appendage-to-body size proportions, such as the pleopod length metric, hold great promise, particularly for species of slipper lobsters like S. squammosus for which there exist no other reliable conventional morphological measures of sexual maturity. Morphometric proportions also should be included among the factors evaluated when assessing size at sexual maturity in spiny lobster stocks; previously, these proportions have been obtained routinely only for brachyuran crabs within the Crustacea.

The survey of sexual maturity in Mugil cephalus so measuring sex hormones and histologic studies

In this study the process of female gray mullet brooders was carried out by using histological study and masurment of sex steroids. Results of histological studies showed that oocyte of gray mullet brooders in Gomishan Rearing Center conditions of develop to the end of yolk globule stage. The results were observed with oocyte in chromatin nucleolar stage (first stage) with means of diameter of 20 p m, in August, perinucleolar stage (second stage) in September with mean diameter of 87 p m, yolk vesicle stage (third stage) in October with mean diameter 200 p m and yolk granules stage (forth stage) from October to November with average diameter of 180 — 650 p m. For the reason of stopping oocyte develop at the end of fourth stage, hormonal induction to final oocyte maturation and ovulation was used. For this purpose, carp pituitary , HCG and LRH-A2 with different combinations were used in two stages, second injection was used 24 hours after first injection. 15 females brooders were divided in 5 groups, different hormonal combinations were injected to four groups and to fifth group as control, only saline, was injected. The process of female brooder rippening in hormonal induction was studied via masurment of sex steroids including 17 a - hydroxy progestrone, estradio1-17)6 and testosterone. Blood samples were collected from caudal vein during first injection, 24, 30 and 48 hours after the first injection. At the same time, for distinguishing histological changes the sample has been attained from the gonads Sex stroid fluctuation patterns in different brooder groups that injected hormon were similar, however hormonal composition had similar effects. All brooder that their oocyte in the beginning of hormonal injection were At the end of fourth stage with oocyte diameter average of 600 p m received to final maturation and ovulation. The brooder that its oocytes were At the begining or mid-fourth stage did not show ovulation but hormonal induction caused oocyte develop at the beginning of fifth stage. Study of 17-hydroxy progestrone fluctuation showed that the maximum level of this steroid (0.347 ng/ml) measured 30 hours after the first injection and was significantly higher (p< 0.05) than those of control group. So, 17-hydroxy progestrone is probably precursor of maturation inducing steroid (MIS). However the maximum level of that observed was coincident with germinal vesicle breakdown, oil droplets coalescence and dissolution of yolk granuls The maximum levels of esteradiol— 17/0 and testosterone (3.778 and 16.801ng/ml,respectively) in spawned brooders,were observed 24 hours after the first injection. levels of those steroids were significantly higher (p<0.05) than control group. Maximum level of sex steroids in the brooders that did not spawn to the end of treatment was observed with more delay than those in spawned brooders. Therefor maximum level of 17a-hydroxy progestrone (0.264 ng/ml) in those brooders observed in fourth sampling time and the maximum levels of estradio1-17a and testosterone (2.944 and 18.993 ng/ml, respectivly)observed in third sampling time that was significantly higher (p<0.05) than those of control group. For the study of stress effect on brooders during the hormonal induction, level of cortisol was measured in every sampling time. level of cortisol had high fluctuation that showed handling level and stress effect on brooders. However maximum level of cortisol in majority of brooders was dominant in third sampling time that was coincident with final maturation.

Seasonal foods, gonadal maturation, and length-weight relationships for nine fishes commonly captured by shrimp trawl on the Northwest Gulf of Mexico Continental Shelf

Recent emphasis on ecosystem approaches to fisheries management renews interest in, and the need for, trophic information about fish communities. A program was started in 1980 at the National Marine Fisheries Service Galveston Laboratory to develop a trophic database for continental shelf fishes. Collections were made during 1982-1983 that were processed but never published, yet the data remain valid today for historical purposes and for delimiting food web components within ecosystem assessments. I examined spring, summer, and fall foods in offshore populations of nine common species of trawl-susceptible fishes, with particular reference to predation on commercial penaeid shrimps (Farfantepenaeus and Litopenaeus). Diets were evaluated with the Index of Relative Importance (IRI) which combines the occurrence, number, and weight of each food item. Bank sea bass (Centropristis ocyurus) and bighead searobin (Prionotus tribulus) primarily consumed crabs, more so by larger than smaller fish. Inshore lizardfish (Synodus foetens) was almost entirely piscivorous. Ocellated flounder (Ancylopsetta ommata) consumed fishes, crabs, and stomatopods. Dwarf sand perch (Diplectrum bivittatum), blackwing searobin (Prionotus rubio), rock sea bass (Centropristis philadelphica), southern kingfish (Menticirrhus americanus), and red snapper (Lutjanus campechanus) fed mainly on shrimps. Most fish diets varied with respect to size (age), time of day, area sampled, depth, or season. Rimapenaeus and Sicyonia were the most frequently identified shrimp genera - only five Farfantepenaeus and no Litopenaeus were identified in almost 4,300 fish stomachs. I also examined gonadal development and documented fish length-weight relationships. Ripe gonads were most frequently found during summer in dwarf sand perch, during fall in ocellated flounder and bighead searobin, and during spring for other species, except no ripe red snapper or bank sea bass were collected. Rock sea bass was found to be a protogynous hermaphrodite, while dwarf sand perch is a synchronous hermaphrodite. Only ocellated flounder and southern kingfish exhibited sex-related differences in length-weight relationships. (PDF contains 40 pages.)

Studies of the age, growth, sexual maturity and spawning of populations of anchoveta (Cetengraulis mysticetus) of the coast of the Eastern Tropical Pacific Ocean

ENGLISH: Three hundred and twenty-six collections of anchoveta (Cetengraulis mysticetus), an important tuna bait species, taken between April 1951 and April 1960 from seven major baiting areas in the Eastern Tropical Pacific Ocean (Almejas Bay, Guaymas, Ahome Point, Banderas Bay, Gulf of Fonseca, coast of Colombia and Ecuador-Peru) are the basis of this study of age, growth, sexual maturity and spawning. The study of the temporal progression of modal size groups from plots of monthly length-frequency distributions provided estimates of age and rate of growth. The study of sexual maturity and time of spawning was based on gross examination of ovaries, and application of the gonad index. SPANISH: Trescientas veintiseis recolecciones de anchovetas (Cetengraulis mysticetus), una importante especie de carnada para la pesca del atún, cogidas entre abril de 1951 y abril de 1960 en siete de las mayores áreas de pesca de peces de carnada en el Océano Pacífico Oriental Tropical (Bahía de Almejas, Guaymas, Punta Ahome, Bahía Banderas, Golfo de Fonseca, y las costas de Colombia y de Ecuador- Perú), sirven de base a este estudio de la edad, crecimiento, madurez sexual y desove de dicha especie. El estudio de la progresión temporal de los grupos de tamaños modales según los gráficos de las distribuciones de la frecuencia de las longitudes proporcionó estimaciones de la edad y de la tasa de crecimiento. La investigación de la madurez sexual y la época de desove se basó en el examen macroscópico de los ovarios y en la aplicación del índice de gónadas.

Sexual Reproduction and Early Development in the Estuarine Sea Anemone, Nematostella vectensis Stephenson, 1935

This dissertation: 1) determines the factor(s) responsible for spawning induction in NematosteJla vectensis; 2) isolates, describes, and documents the source of jelly from egg masses of N. vectensis; and 3) describes N. vectensis' early development. Namatostella vectensis were maintained on a 7-day mussel feeding/water change regime over 159 days. Within 36 hours of mussel feeding/water change. 69.1% of females and 78.5% of males spawned reliably. Through manipulation of feeding, water change, oxygen and nitrogenous waste concentrations, spawning induction was found to be triggered by the oxygen concentration associated with water change, and not by feeding. Ammonia, anemones' major waste product, inhibited this induction in a concentration-dependent manner. Female N. vectensis release eggs in a persistent jellied egg mass which is unique among the Actiniaria. The major component of this egg mass jelly was a positive periodic acid-Schiffs staining, 39.5-40.5 kD glycoprotein. Antibodies developed in rabbits against this glycoprotein bound to jelly of intact egg masses and to granules (~ 2.8 IJm in diameter) present in female anemone mesenteries and their associated filaments. Antibodies did not label male tissues. Nematostella vecfensis embryos underwent first karyokinesis -60 minutes following the addition of sperm to eggs. Second nuclear division took place, followed by first cleavage, 90-120 minutes later. Each of the 4 blastomeres that resulted from first cleavage contained a single nucleus. Arrangement of these blastomeres ranged from radial to pseudospiral. Embryonic development was both asynchronous and holoblastic. Following formation of the 4-cell stage, 71% of embryos proceeded to cleave again to form an 8-cell stage. In each of the remaining 29% of embryos, a fusion of from 2-4 blastomeres resulted in 4 possible patterns which had no affect on either cleavage interval timing or subsequent development. The fusion event was not due to ooplasmic segregation. Blastomeres isolated from 4-celled embryos were regulative and developed into normal planula larvae and juvenile anemones that were 1/4 the size of those that developed from intact 4-celled embryos. Embryos exhibiting the fusion phenomenon were examined at the fine structural level. The fusion phenomenon resulted in formation of a secondary syncytium and was not a mere compaction of blastomeres.

Reproduction, maturation, and seed production of cultured species: Proceedings of the Twelfth U.S.-Japan Meeting on Aquaculture, Baton Rouge, Louisiana, October 25-29, 1983

The United States and Japanese counterpart panels on aquaculture were formed in 1969 under the United States-Japan Cooperative Program in Natural Resources (UJNR). The panels currently include specialists drawn from the federal departments most concerned with aquaculture. Charged with exploring and developing bilateral cooperation, the panels have focused their efforts on exchanging information related to aquaculture which could be of benefit to both countries. The UJNR was begun during the Third Cabinet-Level Meeting of the Joint United States-Japan Committee on Trade and Economic Affairs in January 1964. In addition to aquaculture, current subjects in the program include desalination of seawater, toxic microorganisms, air pollution, energy, forage crops, national park management, mycoplasmosis, wind and seismic effects, protein resources, forestry, and several joint panels and committees in marine resources research, development, and utilization. Accomplishments include: Increased communication and cooperation among technical specialists; exchanges of information, data, and research findings; annual meetings of the panels, a policy-coordinative body; administrative staff meetings; exchanges of equipment, materials, and samples; several major technical conferences; and beneficial effects on international relations. (PDF file contains 79 pages.)

Size composition, growth and sexual maturity of bigeye tuna, Thunnus obesus (Lowe), from the Japanese long-line fishery in the Eastern Pacific Ocean

Size composition data of bigeye tuna taken from the eastern tropical Pacific Ocean by Japanese Prefectural experimental training vessels from 1958 to 1964 are examined. A gradient of increasing fish size from east to west is noted. Males increase in ratio over females for the entire range of lengths examined, and beyond 170 cm comprise more than 75 per cent of the total. The first semester of the year is important as a bigeye spawning season. A general relationship between sexual maturity and thermal structure of the water is discussed. At the end of their 12th quarter of life bigeye are about 114 cm long, by the 16th quarter, 137 cm and at the end of 20 quarters, about 153 cm. The long-line fishery in the eastern Pacific has had a marked effect on the size composition of the stocks of bigeye, but whether the fishing has driven the stocks below a point which could afford a maximum sustainable yield could not be determined. (PDF contains 55 pages.)

Studies of the sexual development and spawning of yellowfin tuna, Neothunnus macropterus, and skipjack, Katsuwonus pelamis, in three areas of the Eastern Pacific Ocean, by examination of gonads.

ENGLISH: Knowledge of the size and age at maturity, spawning seasons, and spawning areas of the tropical tunas supporting the fishery in the Eastern Pacific is an important part of the basic information required for understanding their life history, population structure, and fishery dynamics. Until a few years ago nothing was known of these matters. In 1947 the senior author and one of his colleagues (Schaefer and Marr 1948, Schaefer 1948) were able to demonstrate that both yellowfin tuna and skipjack spawn offshore from Central America at least during the late winter and spring months. During January to April many yellowfin tuna over about 70 cm. total length in commercial catches from that region were found to have gonads in advanced stages of maturity, and specimens caught during late June were found to be spent. Maturing skipjack were collected in late February, and spawned-out fish were observed in late March. Numerous very young juveniles of the yellowfin, down to 10 mm. in length, and two very young juvenile skipjack, were captured in this area between January and May. SPANISH: El conocimiento del tamaño y la edad que corresponden a la primera madurez sexual, así como de las estaciones y áreas de desove de los atunes tropicales que mantienen las pesquerías del Pacífico Oriental, constituyen parte importante de la información que es menester para comprender la historia natural, la estructura de la población y la dinámica de la pesquería. Hasta hace pocos años nada se sabía sobre el particular. En 1947 el autor principal y uno de sus colegas (Schaefer y Marr, 1948; Schaefer, 1948) pudieron demostrar que tanto el atún aleta amarilla como el barrilete desovan en el mar abierto, frente a América Central, por lo menos durante la última parte del invierno y en la primavera. De enero a abril encontraron que muchos de los atunes aleta amarilla de más de 70 cm. de longitud total, procedentes de las pescas comerciales de dicha región; tenían gónadas en avanzados estados de madurez, mientras que ejemplares pescados hacia fines de junio ya habían desovado. Se recolectaron barriletes en vías de maduración a fines de febrero, al paso que en los últimos días de marzo se encontraron especímenes que ya habían desovado. Numerosos ejemplares muy juveniles del atún aleta amarilla, tan pequeños como 10 mm., y dos barriletes también muy juveniles, fueron pescados en esta región entre enero y mayo. (PDF contains 65 pages.)

A study of the age, growth, sexual maturity, and spawning of the anchoveta, Cetengraulis mysticetus, in the Gulf of Panama

ENGLISH: Crew members of tuna clippers and Commission personnel are collecting specimens of anchovetas (Cetengraulis mysticetus) for studies of the biology of this important tuna-bait species. More than 27,000 fish from 231 collections captured in the Gulf of Panama between June 1951 and January 1956 are the basis of this study of the age, growth, sexual maturity, and spawning season of this species in that area. Estimates of age and rate of growth were made by studying the temporal progression of modal size groups from monthly length frequency distributions. Sexual development and time of spawning were determined from gross examination of ovaries and measurements of ovarian eggs. SPANISH: Con el fin de estudiar la biología de la anchoveta (Cetengraulis mysticetus) los tripulantes de los barcos atuneros y el personal de la Comisión están recolectando especimenes de esta importante especie de carnada para capturar el atún. Mas de 27,000 ejemplares de las 231 colecciones hechas en el Golfo de Panamá entre junio de 1951 y enero de 1956, sirven de material al presente estudio sobre la edad, el crecimiento, la madurez sexual y las épocas de desove de esta especie en el área indicada. Las estimaciones de la edad y de la proporción del crecimiento fueron hechas a base del estudio de la progresión temporal de los grupos modales de tamaño en las distribuciones mensuales de frecuencias de longitud. El desarrollo sexual y el periodo de desove fueron determinados mediante el examen microscópico de los ovarios y las mediciones de los huevos ováricos. (PDF contains 79 pages.)

Sexual maturity, fecundity and egg size of wild and cultured samples of Bagrus bayad macropterus

Twenty four matured samples of Bagrus bayad macropterus from the wild (Shiroro Lake, Nigeria) and under captivity, size ranging from 412.69-3300.00 g total body weight, were analysed for sexual maturity,fecundity and egg size. The average fecundity obtained were 53352.59 and 21028.32 eggs for the wild and cultured fish respectively.Positive relationship was observed between fecundity, body size and gonad weight. Fecundity increased as body size increased. A more positive and linear relationship was observed between fecundity and gonad weight than fecundity and total body weight. Egg diameter,length and weight were determined from the egg samples. The mean size range of eggs for cultured fish was 0.74-1.05 mm of diameter; 1.01-1.20 mm of length and 0.25-0.40 mg of weight. Wild samples had mean size range of 0.68-l.09 mm of diameter, 0.85-1.38 mm of length and egg mean weight range was 0.15- 0.40 mg. Sexual maturity is dependent on size (1 kg and above). The egg diameter, egg length and weight bear no relationship with each other. Gonad development study indicated that gonad development was faster under captivity than in wild

Environmental oestrogens and sexual development in fish

This brief paper discusses the assumption that watercourses might be harbouring a chemical(s) affecting the sexual development in fish. Male fish was found with the oestrogen-dependent blood protein, vitellogenin, usually found only in maturing females. The author examines a number of man-made chemicals present in the environment have been found to be oestrogenic. The paper concludes that rivers contain environmental oestrogens that are capable of causing disruptions in the sexual development of fish. Whether or not these environmental oestrogens are causing a widespread disruption in reproduction in wild fish, however, has yet to be determined.

Modulation of the effectiveness of 17-alpha-hydroxy-20-beta-dihydroprogesterone or of a gonadotrophic extract on the in vitro intrafollicular maturation of oocytes of the rainbow trout Salmo gairdnerii by various non-maturing steroids [Translation from: Compte Rendu Hebdomadaire des Seances de l'Academie des Sciences, Paris, Series D 281, 811-814, 1975]

The effectiveness of 17 α-hydroxy-20 β-dihydroprogesterone (17 α-20 β Pg) or of a trout hypophyseal gonadotrophic extract on the in vitro intrafollicular maturation of trout oocytes can be modulated by steroids which do not have a direct maturing effect; the effectiveness of the gonadotrophic extract is lowered by oestradiol and oestrone and increased by testosterone. As these steroids have no significant effect on maturation induced by 17 α-20 β Pg, the site of their activity is probably in the follicular envelopes. Corticosteroids, and Cortisol and cortisone in particular increase the effectiveness of the gonadotrophic extract, but increase the effectiveness of 17 α-20 β Pg even more strongly, suggesting that this 'progestagen' has a direct effect on oocyte sensitivity.