The study of biological characteristics and heavy metal determination of Cd, Pb, Fe, V, Ni in the muscle, liver and gonad tissues of kingfish Scomberomorus commerson in the coastal waters of the Hormozagan Province (Persian Gulf)

Biological studies and heavy metal (Ni, v, Fe, Pb,Cd) determination in liver, gonad and muscle of Scomberomorus commerson were carried out from Oct 2006 to Sept 2007 in Hormozgan coastal waters. 599 Samples were gutted for reproduction and nutrition studies, fork length and weight were measured to nearest cm and g respectively. Meanwhile 40 samples were also investigated for heavy metal studies. All specimens were collected from two major landing sites (Bandar Lengeh & Bandar Abbas). Minimums & maximum fork length & weight were 29, 128 cm & 235 and 15350 g respectively.Isometric growth was shown according to our study and b was estimated 2.9 (overall), 2.91(male) & 2.89 (female). The average relative gut length was 0.52± 0.007 and it was determined that S. commerson is a carnivorous. More than 99 percent of gut content was different teleost fishes. Gastro somatic index had two peaks in Nov & Jan (before spawning) and with a decreased trend in July, the spawning period. Occurrence of empty stomach was estimated % 65.77. Maximum amount of condition factor was in Dec. Spawning season was started from June. The average of Absolute & relative fecundity (to weight unit) was 1217149±179315 and 178.2±15.58 respectively. Lm50% was estimated 75 cm for females. Sex ratio was 0.97: 1 (male: female). Chi- Square test showed no significant difference (p>0.05). Maximum amount of hepatosomatic index was estimated in March.Metal concentrations were determined using either Flame Atomic Absorption Spectroscopy (for Fe) or Graphite Furnace Atomic Absorption Spectroscopy (for Pb,Cd,Ni and V). The mean concentration (μg/g dry weight)of Pb,Cd,Ni,V and Fe in the liver were 0.0309, 0.0268, 0.0672, 0.0077, 2.5159 in the gonad 0.0440 ,0.0295, 0.1096, 0.0000, 1.4449 and in the muscle 0.0244, 0.0324, 0.0656, 0.0128, 1.6138 respectively. The maximum metal concentrations were below the maximum permissible limits for human consumption recommended by the USEPA, WHO and the UK. The results of Kendall's Tau-b correlation coefficient were as follows: The Liver tissue: There were significant positive linear relationships between accumulation of V, Fe, and Pb with Fork length, Pb and Fe with weight, GSI with Pb, Cd, V and 109 Fe, and a negative linear relationships between HSI with accumulation of V and Fe, Fork length, weight and GSI. The Gonad tissue: There were significant positive linear relationships between GSI with accumulation of Pb, Cd, Fe, Fork length and weight, a negative linear relationship between HSI with Fork length, weight and GSI. The Muscle tissue: There were significant positive linear relationships between accumulations of V, Fe with Fork length and weight factors and as well as GSI with Cd, V, Fe, Pb, Fork length and weight,a negative linear relationship between HSI with Fork length, weight, Cd, Fe and GSI. The results of Mann-Whitney U tests (P≤0.05) show that there were significant differences between summer and autumn from heavy metal contents in the studied tissues point of view. The only exceptions were for Ni in the liver, gonads and muscle and as well as there were significant differences between male and female from heavy metal contents in the studied tissues. The only exceptions were for Pb in muscle, Ni in liver, gonad and muscle, V in muscle, and Cd and Fe in gonads.

CORRELATION OF ZONA-BINDING WITH OOCYTE MATURATION AND SPERM MOTILITY IN RHESUS-MONKEYS BY HEMIZONA ASSAY

The hemizona assay (HZA) in Rhesus monkeys was employed to study the correlation of zona-binding ability with sperm motility or with naturally developing oocytes at various maturational stages. Oocytes from unstimulated ovaries were retrieved within 2 hr from monkeys sacrificed for vaccine production (in reproductive season, but with their menstrual cycles not determined). Oocytes were divided into four groups based on their morphological maturation: 1) Oocytes surrounded by more than one cumulus layer (MC); 2) Oocytes retaining intact germinal vesicle nuclei (GV); 3) Oocytes with germinal vesicle breakdown showing distinct perivitelline space (PVS); and 4) Oocytes extruding the first polar body (PB1). The mean numbers of sperm bound to hemizona for PBI, PVS, GV, and MC groups were 132.9 +/- 12.0, 71.5 +/- 10.1, 36.1 +/- 4.0, and 20.1 +/- 2.9 (Mean +/- SE), respectively. The four groups showed significant differences from each other in sperm/egg binding ability (P < 0.01). The number of bound sperm significantly increased with oocyte maturation. The present study also showed that zona-binding ability was also affected by sperm motility. For sperm with 67.7% motility and sperm with 31.2% motility, the average numbers of bound sperm were 43.5 +/- 2.2 and 25.3 +/- 2.9 (Mean +/- SE), respectively. There was significantly higher binding ability for sperm with higher motility (P < 0.01). The results suggest that: 1)The rhesus monkey model can serve as a very sensitive model for studying sperm/egg interaction by HZA; 2) Sperm motility positively correlated with sperm/egg binding; and 3) Sperm/egg binding ability increases with oocyte maturation. The binding ability is highest when oocytes matured to the PB1 stage, which is also the best opportunity for fertilization. This is strong evidence for the ''zona maturation'' hypothesis. (C) 1994 Wiley-Liss, Inc.

Study on some biological and feeding factors of spined loach Cobitis taenia from the Babolrud, Talar and Siahrud Rivers

Spined loach, Cobitis taenia, is a predominant fish in the river systems of the southern Caspian Sea basin. Although there is evidence of the geographical divergence of this taxon, but no information is available on morphological differences within the species populations. This study was designed to evaluate some biological factors including; morphometric and meristic characters, length-weight, age-growth, condition factor, diet, reproduction, variation and differentiation, in the Babolrud, Talar and Siahrud Rivers in south of the Caspian Sea basin. Age, sex ratio, fecundity, ova diameter and gonadosomatic index were estimated. Also, regression analyses was tested the relation between fecundity and fish length, weight, gonad weight, and also age. Totally 858 fish of which 721 were matures, were collected from these rivers by electrofishing. 37 morphometric characters, 9 meristic characters and 78 truss network system characters were estimated. Resulats of DFA analysis based on data of morphometric and meristic showd that these populations are highly (94.5%) varios from each other. In discriminate function analysis, the proportion of individuals correctly classified into their original groups was 61%, 65.4% and 86.5% for upstream and downstream, respectively. Clustering based on Euclidean distances among groups of centroids using an UPGMA and also principal component analysis’ results for morphometric data indicated that these populations from these three rivers were clearly distinct from each other. Regression equations between length and weight in these three populations were significantly different from Folton factor (b=3), that showed the fish has a negative Alometric growth process. Condition factor was estimated between 0.8912 to 1.2736 and 0.8131 to 1.4489 for males and females, respectively. Sex ratio (female: male) in these populations was 1.2816:1. The difference between the number of females and males was significant and females were more than males. The female and male specimens reach maturity by Tl more than 40 and 30 mm and at the age of 2+ and 1+, respectively. The mean of ova diameter was 0.5824±0.2882. The spawning took place from May to late July, at the water temperature from 18.7 to 24.0°C. The GSI values average at the beginning of the reproduction period was about 9%, with ranged from 2 to 26% in ripe mature females. The absolute and relative fecundity were 2109±792 and 579±208 respectively. The absolute fecundity was significantly related to body weight and gonads weight. Based on the pattern of gonado-somatic index, it was concluded that this fish has prolong active reproductive period, which is a type of adaptation by short-lived small fishes to environmental conditions. The macroscopic and histological results showed that the female and male have 5 and 4 stages in their maturation process, respectively. The RLG index was about 0.4732, which showed the fish is a carnivorous species. Significant difference was observed between fishes with different length and diet. The main foods of the fish were Trichoptera, Chironomidae larvae and Ephemeroptera which were their prefered food as well, however it was estimated that the food selection and diet are affected by environmental conditions.

Amino acid requirements for maturation of rhesus monkey (Macacca mulatta) oocytes in culture

This study evaluated the effects of different amino acid formulations on supporting meiotic and cytoplasmic maturation of rhesus monkey (Macacca mulatta) oocytes in vitro. Five hundred and forty-six cumulus-oocyte complexes (COCs) aspirated from unstimulated adult monkey follicles (greater than or equal to 1000 mum in diameter) were cultured in either modified Connaught Medical Research Laboratories 1066 medium (mCMRL-1066) or in one of eight chemically defined media (modified basic medium 5 supplemented with 5.5 mmol glucose l(-1), 0.003 mmol pantothenic acid l(-1) and different amino acid formulations) as below: (1) modified basic medium 5 (mBM5) containing no amino acid; (2) mBM5 + 0.2 mmol glutamine l(-1); (3) mBM5 + 11 amino acids from hamster embryo culture medium 6 (HECM-6) (11 AA); (4) mBM5 + Eagle's non-essential amino acids (NEA); (5) mBM5 + NEA + 0.2 mmol glutamine l(-1); (6) mBM5 + Eagle's essential amino acids (EA) without glutamine; (7) mBM5 + EA + 0.2 mmol glutamine l(-1); (8) mBM5 + Eagle's 20 amino acids (20 AA) + 0.2 mmol glutamine l(-1); and (9) mCMRL-1066 (control). All media contained FSH, LH, oestradiol and progesterone. After maturation, mature oocytes were subjected to the same fertilization and embryo culture procedures. COCs matured in treatment 5 had greater potential to progress to metaphase II (66%; P < 0.05) than did those in treatments 1 (37.3%), 2 (48.3%)f 3 (41%), 6 (41%) and 9 (43%). Oocytes matured in treatment 8 had the best morula (53%) and blastocyst (18%) developmental responses (P<0.05). The lowest (P<0.05) morula and blastocyst developmental responses were obtained from COCs matured in treatments 1 (0%) and 6 (8%). The other media supported intermediate embryonic development (range 11-38% of morula and blastocyst). These results indicate that the choice of amino acids affects the competence of oocyte maturation and that Eagle's 20 AA with 0.2 mmol glutamine l(-1) is more efficient than the other amino acid formulations for maturation of rhesus monkey oocytes.

Investigation on reproduction process in two species of crab, Eriphia sebana and Ocypode saratan in the intertidal zone of Chabahar area

The present study with headline investigation on reproduction in two species of crab Eriphia sebana and Ocypode saratan was carried out in the intertidal zone of Chabahar in thirteen month from December 2004 to December 2005. Checked samplings have been taken, 45 number Crab monthly from any four stations by manual or use trap. During this study the following subjects were measured: temperature range and salinity, measurable coast granule, determination of sex ratio, relations carapace width with carapace length, Carapace width with total body weight, Gonad weight, gonadosomatic index, condition factor, gastrosomatic index, investigation content in stomach, LM50, growth parameters, plenty distribution length and width and gonad weight and total body weight. Studied on measurable coast granule were expressed that Ocypode saratan in Desalination station, were nest in soils equable sand and this quantity were confirmed in Pozm station. Sex ratio were assign in desalination area and Pozm M: F 0/44:0/56 and in Tiss and Chabahar M:F 0/45:0/55. Carapace length and carapace width (cm) and body weight (g) Furthest were designated in Ocypode saratan within carapace width sequential: In female: 5/42-6/15-105/13 and in male: 5/53-6/25-108/91 and in Eriphia sebana within Tiss area sequential: in female: 5/12-5/94-110/21 and in male 5/14-60/01-114/37. Have been linear relationship between carapace length and carapace width and equaled CW=aCL+b. Weight growth in two species were be modal and equaled BW=aCLb And increased crab weight by built up carapace width. Maximum gonad weight in Ocypode saratan within Desalination area in female have been outcome 3/39 and in male 0/84g and in Eriphia sebana extreme within Tiss during may in female were be 4/18 and in male 1/1g. Stomach content in Eriphia sebana were involved a black until half-purplish liquid and yellowish in Ocypode saratan. Stomach contents identifiable were being in four groups: Molluscoid, Crustacean, Plankton and Fish. Carapace width during the first year of maturation have been LM50:3/77 in Desalination area and LM50:3/92 in Pozm for Ocypode saratan and LM50:4/26 in Tiss and LM50:4/62 in Chabahar. Ability spawning in Eriphia sebana within Tiss has been CW=4/17cm and in Ocypode saratan within Desalination area CW=4/23cm. Maximun value of Loo for Eriphia sebana was equal 59/67 and growth factor K=0/68 within Tiss and Loo =61/64, K=0/65 for Ocypode saratan within Desalination area. Maximun GSI and GI have been within Desalination area and Tiss and minimum within Pozm and Chabahar. The maturity stages of two species were classifed into six stages. Review on GSI, CF have been showed that relation with temperature and salinity and definer in two species have been spawned in two period that Maximun in spring premier than autumn.

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.

Study on the reproductive biology of Pinctada fucata (Gould) (Mollusca: Pteriidae)

Annual cycle of gonad development and spawning in pearl oyster, Pinctada ficata (Gould) in Nakhiloo, Northeast Persian Gulf, was investigated over two years from August 1994 to June 1996. Gonadal condition was assessed by staging criteria to describe gametogenic development from histological preparations of randomly collected individuals of all sizes. A bimodal gametogenic pattern with summer and autumn spawning periods was evident throughout the study. Gametogensis commenced in November-December which proceeded by major gonadal maturation during February-April. Summer spawning was observed from April to July with major spawning at the latter end. During spawning peak in July, low level of gametogensis was noticed. Gametogenic activity was picked up again in August-September which proceeded by autumn spawning from September to December. Towards the end of spawning season, incidence of gonadal inactivation increased. Minimum level of gonadal activity was observed in November. Temperature regime appears to have influential role in regulation of gametogenic and spawning processes. Gonadal development and spawning trends were similar in both sexes. P. radiaata was found to be protandrous hermaphrodite which matured as a male at shell height greater than 20 mm. Biseivality was uncommon and the sex ratio was about 1:1. Ultrastructure of gametes were investigated in the Pictada fucata (Gould). "Auxiliary cells" closely accociated with developing oocytes were observed. Each oocyte seems to be associated with only one secretory cell. which is characterized by an abundant rough endoplasmic reticulum at the onset of vitellogenesis. Contact between this cell and a developing oocytes is maintained by a desmosome-like junction which can be observed when the vitelline coat is formed. these "auxiliary or nursing cells" seem to play a tropic role in vitellogenesis, and may be involved in the formation of the vitelline coat of the oocytes. Oocytic degeneration is observed in this species, it is a continuous phenomenon of varing intensity throughout the year. The ultrastructural changes resulting in lysis of the oocyte are described. Mature spermatozoa consist of a broad, cap-shaped acrosomal vesicle, subacrosomal material, a round nucleus, two triplet substructure centrioles surrounded by four spherical mitochondria, and a flagellum anchored to the distal centriole and plasma membrane. Spermatozoa of Plucata closley resemble to those of other investigated Pteriidae. Changes in proximate composition of soft tissue and gonadal cycle of Pinctada fucata was studied. Mobilization and utilization of stored reserves are apparent during gametogenesis and gonadal maturation. Protein reserves are utilized during spermatogenesis while reserved carbohydrates form the main energy donor in oogenesis. The role of lipid as am.: energy reserve is second to that of carbohydrate.

La reproducción de la merluza en el Mar Argentino (Merlucciidae, Meluccius merluccius hubbsi) : 1. Descripción histológica del ciclo del ovario de merluza

The present paper deals on the histological description of the hake ovary made on the basis of gonad observations of 394 females during the period April 1966 March 1967. The material was obtained from weekly sampling of commercial catches carried out at the Institute of Marine Biology (Prov. Buenos Aires, Argentina). The anatomical and histological description of the standard ovary and the adopted terminology are given. The maturation process is divided into five periods, from ovocyte formation to yolked ovocyte formation, with its histological description. Ovary changes are analyzed on detail. The following conclusions were outlined: 1. Analysis demonstrated that although some specimens were totally spawned others, at the end of the spawning period, retaining a great number of ovocytes in different maturity stages. Therefore, postspawners have been classified as follows: Postspawned II : This stages is characterized by the empty ovarian structure, with ovocytes in stage II, which will remain in the resting phase untill next spawning season. Postspawned III and IV: Their main characteristics are: tissue destruction, bloody residuals and remaining ovocytes in stage III and IV, respectively. 2. Some transformations were found in ovaries of postspawned III and IV. They are classified as follows, according to its origin and structure: Developed from follicular cell membrane – a) Glandular formations, b) Epiteloid formations - Originated from remaining ovocytes, c) Ovocyte disintegration, d) Ovocyte with follicular cell infiltration. 3. All those structures derived from postpawners III and IV have a temporary character and will be reabsorbed. Their presence delay the recuperation of the organ and its reproductive functions. Consequently, the possibility of those structures acting as control mechanisms is suggested. 4. Transformations pointed out in paragraph Nº 2 prevent the possibility of consecutive spawning originated from the remaining ovocytes (II and IV). 5. No structures originated from postspawners III and IV were found during summer season. 6. Reproductive cycle of hake has been described monthly. It was observed that maturing ovaries predominate in summer (November-December).

Nanobodies raised against monomeric α-synuclein distinguish between fibrils at different maturation stages.

Nanobodies are single-domain fragments of camelid antibodies that are emerging as versatile tools in biotechnology. We describe here the interactions of a specific nanobody, NbSyn87, with the monomeric and fibrillar forms of α-synuclein (αSyn), a 140-residue protein whose aggregation is associated with Parkinson's disease. We have characterized these interactions using a range of biophysical techniques, including nuclear magnetic resonance and circular dichroism spectroscopy, isothermal titration calorimetry and quartz crystal microbalance measurements. In addition, we have compared the results with those that we have reported previously for a different nanobody, NbSyn2, also raised against monomeric αSyn. This comparison indicates that NbSyn87 and NbSyn2 bind with nanomolar affinity to distinctive epitopes within the C-terminal domain of soluble αSyn, comprising approximately amino acids 118-131 and 137-140, respectively. The calorimetric and quartz crystal microbalance data indicate that the epitopes of both nanobodies are still accessible when αSyn converts into its fibrillar structure. The apparent affinities and other thermodynamic parameters defining the binding between the nanobody and the fibrils, however, vary significantly with the length of time that the process of fibril formation has been allowed to progress and with the conditions under which formation occurs, indicating that the environment of the C-terminal domain of αSyn changes as fibril assembly takes place. These results demonstrate that nanobodies are able to target forms of potentially pathogenic aggregates that differ from each other in relatively minor details of their structure, such as those associated with fibril maturation.

Nanobodies raised against monomeric α-synuclein distinguish between fibrils at different maturation stages

Nanobodies are single-domain fragments of camelid antibodies that are emerging as versatile tools in biotechnology. We describe here the interactions of a specific nanobody, NbSyn87, with the monomeric and fibrillar forms of α-synuclein (αSyn), a 140-residue protein whose aggregation is associated with Parkinson's disease. We have characterized these interactions using a range of biophysical techniques, including nuclear magnetic resonance and circular dichroism spectroscopy, isothermal titration calorimetry and quartz crystal microbalance measurements. In addition, we have compared the results with those that we have reported previously for a different nanobody, NbSyn2, also raised against monomeric αSyn. This comparison indicates that NbSyn87 and NbSyn2 bind with nanomolar affinity to distinctive epitopes within the C-terminal domain of soluble αSyn, comprising approximately amino acids 118-131 and 137-140, respectively. The calorimetric and quartz crystal microbalance data indicate that the epitopes of both nanobodies are still accessible when αSyn converts into its fibrillar structure. The apparent affinities and other thermodynamic parameters defining the binding between the nanobody and the fibrils, however, vary significantly with the length of time that the process of fibril formation has been allowed to progress and with the conditions under which formation occurs, indicating that the environment of the C-terminal domain of αSyn changes as fibril assembly takes place. These results demonstrate that nanobodies are able to target forms of potentially pathogenic aggregates that differ from each other in relatively minor details of their structure, such as those associated with fibril maturation. © 2013 Elsevier Ltd.

Distribution of microcystins in various organs (heart, liver, intestine, gonad, brain, kidney and lung) of Wistar rat via intravenous injection

The distribution of microcystins (MCs) in various tissues of Wistar rats was studied under laboratory conditions. Rats were injected intravenously (i.v.) with extracted MCs at a dose of 80 mu g MC-LRequivalent/kg body weight. MCs concentrations in various tissues were detected at 1, 2. 4, 6, 12 and 24 h post-injection using liquid chromatography-mass spectrometry (LC-MS). The highest concentration of MCs was found in kidney (0.034-0.295 mu g/g dry weight), followed by lung (0.007-0.067 mu g/g dry weight), stomach (0.010-0.058 mu g/g dry weight) and liver (0.003-0.052 mu g/g dry weight). The maximum MCs content in the whole body of rat, 2.9% of the injected dose, was observed at 2 h post-injection. MCs concentration was higher in kidney than in liver during the experiment, and two peaks of MCs concentration (at 2 and 24 h, respectively) were observed in kidney, indicating that MCs can be excreted directly via kidney of rat. Though heart, intestine, spleen, brain, gonad and stomach contained less than 0.2% of injected MCs during the whole experiment stage, the presence of MCs in these tissues represents potential damage to them. (c) 2008 Elsevier Ltd. All Fights reserved.

Gonad development of an anadromous fish Coilia ectenes (Engraulidae) in lower reach of Yangtze River, China

Anadromous Coilia ectenes was sampled from the Yangtze estuary at Chongming and two of the primary upstream spawning grounds at Jingjiang and Anqing in April, May, June and August 2006. Gonad development was analyzed for females. In April, fish were collected in the estuary and at Jingjiang, but not at Anqing. No female was mature (gonad at stages IV or V) at either location. In May, 45% of the females were mature in the estuary, 9% at Jingjiang and 5% at Anqing. In June, 86% were mature in the estuary, 83% at Jingjiang and 7% at Anqing. In August, C. ectenes was absent at Jingjiang. No female was mature in the estuary, and all females were mature at Anqing. Absolute fecundity (AF) increased significantly with standard length (SL) by a power function AF = 2.27 x 10(-6) x SL2.67 (r(2) = 0.57, n = 48, P < 0.05). Mature females in the estuary were smaller than those at Jingjiang and Anqing. Conservation of spawners in the upstream spawning grounds is important because they have a size-related fecundity advantage over the smaller spawners in the estuary.