Reproductive patterns within sub-populations of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

The annual reproductive cycle of the sea urchin Lytechinus variegatus (Lamarck) was examined in individuals collected from four habitats in the Anclote estuary near Tarpon Springs, Florida. Two periods of advanced gametogenic activity were apparent, but this bimodal pattern was not necessarily reflected by gonad growth curves. Differences in reproductive events among sub-populations were related primarily to patterns of gonad growth. Reproductive synchrony present during spring became less apparent as spawning and recovery within the sub-populations took place at differential rates during late-spring and summer. The final spawnout in late summer returned the sub-populations into gametogenic synchrony.     

Factors leading to cannibalism in Lytechinus variegatus (Echinodermata: Echinoidia) held in intensive culture

Ecologically, sea urchins are an important species in marine habitats around the world. Economically, sea urchins are prized for their gonads (uni). With fisheries declining worldwide, intensive sea urchin culture has been proposed. For urchins in commercial culture, any factor that affects survivorship negatively should be addressed to maximize cost benefit. One potential obstacle to optimizing culture of sea urchins is cannibalism. Approximately 2000 adult and juvenile Lytechinus variegatus (1 g-45 g) were collected from Port Saint Joseph Peninsula State Park, FL between June and September 2009. Urchins were held in recirculating tanks at different sizes, densities, and feeding regimes for 4 weeks. Starvation and high density contributed to the highest level of cannibalism among small (12-21 g) urchins (percent cannibalism = 18.8%), whereas fed, high density conditions contributed to the highest level of cannibalism among large (32-37 g) urchins (percent cannibalism = 18.4%). These results suggest that (1) small urchins cannibalize at higher rates than large urchins, and (2) increased density is an important contributing factor leading to cannibalism. We quantified stress, defined as a decrease in production as a result of environmental conditions, by evaluating weight gain within each treatment and suggest that weight loss or minimal weight gain is an indicator of stress. We hypothesize increased stress caused by competitive interference can lead to increased cannibalism and decreased growth rates, even when food is not limiting. Ecologically, there are no reports of cannibalism of urchins in wild populations. Consequently, the role of cannibalism in regulating sea urchin community structure is not known. However, factors affecting cannibalism of L. variegatus in the laboratory may provide insight into the conditions that could result in cannibalism in wild populations. From an aquaculture perspective, it is important to determine those factors that contribute to the incidence of cannibalism in sea urchins so that the appropriate culture conditions can be maintained to reduce the incidence of cannibalism.     

Phagostimulation of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

Artificial agarose foods containing either starch, glycogen, carragheenan, alginate, casein, gelatin, albumin, or peptones at a concentration of 1% dry weight were not phagostimulatory. Artificial foods containing sugars, especially galactose, at a concentration of 0.005 M were phagostimulatory. Of the common amino acids, only L‐phenylalanine was phagostimulatory at this concentration. The initiation of feeding postures (movement of the spines, tube feet, peristome, and Aristotle's lantern) of Lytechinus variegatus was little affected by the concentration of plant extracts. Many aspects of this behavior were attributable to physical rather than chemical stimulation. Feeding rates of L. variegatus upon artificial agarose foods containing either Thalassia testudinum or Gracilaria verrucosa were directly dependent upon plant tissue concentration. All artificial agarose food‐items were initially moved to the mouth and fed upon, however the time spent feeding was directly dependent upon plant type and tissue concentration. Retention of the artificial food‐items upon the test after feeding was not influenced by plant type or tissue concentration. Differences in consumption of foods by L. variegatus are determined by the degree of persistence in feeding upon encountered items and not be selection before initiating feeding or by retention of food‐items for subsequent feedings.     

The effect of temperature-salinity combinations on the functional well-being of adult Lytechinus variegatus (Lamarck) (Echlnodermata,Echinoldea)

The activity coefficient (1000/righting time in sec) was measured to indicate the functional state of Lytechinus variegatus (Lamarck) after exposure to combinations of temperature (22°, 28°, and 34°C) and salinity (25, 30, 35, and 40 ‰). Environmental levels of these variables were 30–33°C and 34–35 ‰. The results indicate that the species lives closer to the upper than lower lethal limits of temperature and salinity. The maximal activity coefficient (18 ± 8) was at 28°C and 35 ‰. A reduction in salinity was probably responsible for a recent mass mortality of the echinoid reported in the eastern Gulf of Mexico.     

The effect of the quality of food patches on larval vertical distribution of the sea urchins Lytechinus variegatus (Lamarck) and Strongylocentrotus droebachiensis (Mueller)

Although most invertebrate larvae are weak swimmers and act as passive particles on horizontal scales, they may be able to regulate their vertical position in response to different factors, including increased food concentration. We examined the effect of the quality of food patches on larval vertical distribution for the sea urchins Lytechinus variegatus and Strongylocentrotus droebachiensis, and determined the effect of dietary conditioning on that response in the laboratory. We reared larvae on a mixed algal diet of Dunaliella tertiolecta and Isochrysis galbana under low (500 cells ml⁻¹) and high (5000 cells ml⁻¹) rations. Food patches were maintained in Plexiglas rectangular columns (30×10×10 cm) using a density gradient, where practical salinity in the bottom layer was 33, in the middle layer 30, and in the top layer 27. We examined the magnitude and mechanism of a behavioural response of larvae of L. variegatus in the four-arm stage, and on two developmental stages of S. droebachiensis (four- and six-arm), by manipulating patch quality. In the absence of a patch, larvae of both species and developmental stages swam through to the surface of the experimental columns. In the presence of algae, fewer larvae were present above the patch and more were at the patch than in control columns. More larvae swam through patches of “unflavoured” algal mimics than algal patches, and aggregated at the surface. Larval distribution relative to patches of algal filtrate without algal cells or of “flavoured” algal mimics in algal filtrate was not consistently different from that in either control or algal patches; thus, the magnitude of larval response to filtrate (with or without particles) was intermediate between that to control and algal patches. For L. variegatus, more larvae crossed the patches when reared on low than high rations, indicating that poorly conditioned larvae may be less responsive to environmental cues. Our results suggest that larvae can actively aggregate and maintain a vertical position in response to a food patch that depends on the quality and quantity of food. The response appears to be based mostly on a chemosensory rather than a mechanosensory mechanism.     

Phylogeography of the sea urchin Paracentrotus lividus (Lamarck) (Echinodermata:Echinoidea): first insights from the South Tyrrhenian Sea

The sea urchin Paracentrotus lividus (Lamarck) (Echinodermata: Echinoidea) is an Atlanto-Mediterranean species abundant in the littoral zone, where it occurs in the sublittoral down to 20 m. The aim of our work is to investigate the genetic patterns of P. lividus along the South Tyrrhenian coasts. Five specimens were collected in six localities, from the Gulf of Naples and the Cilento coast. The nuclear rDNA ITS2 spacer and the two mitochondrial genes 16S and COI were used for the analysis. The three markers utilised did not show any structure among populations from the Gulf of Naples. All populations appear to be polyphyletic, with Cilento samples more differentiated from the others. This suggests the existence of phylogeographic structure at larger geographic scale. Absence of genetic structure has been interpreted taking into consideration theoretical dispersal of the planktotrophic larvae, which can survive for 4–8 weeks before settlement, marine current patterns and persistence of the species in the area.     

Echinoids (Echinodermata: Echinoidea) from the Gulf of Mexico

The echinoid fauna of the Gulf of Mexico collected during three research cruises (20-1260 m depth) was surveyed from samples were taken at 43 stations. A total of 190 individuals were identified (eight orders, 11 families, 15 genera and 18 species). Six species are new records for the Gulf of Mexico: Stylocidaris lineata, Phormosoma placenta placenta, Plesiodiadema antillarum, Plethotaenia spatangoides, Brissopsis atlantica and Hypselaster limicolus. This adds to the little information available on the echinoid fauna of Tamaulipas, Veracruz, Tabasco, Campeche and Yucatan states in Mexico.     

Potential for paired vestibules in plutei (Echinodermata, Echinoidea)

Abstract. Echinoids are generally thought to develop only a single ectodermal invagination (V1), which expands into a vestibule and contributes to the formation of the juvenile rudiment on the left side of the larval body. However, in 2 echinoid species, Strongylocentrotus purpuratus and Dendraster excentricus, a second invagination (V2) was observed to form on the right side of the larval body exactly opposite V1. Formation of the invaginations was followed and characterized using light microscopy and antibodies directed against the S2amide echinoderm neuropeptide as a morphological tag. V2 consistently formed later than V1, and appeared capable of contributing to the formation of a second rudiment when it received the appropriate mesodermal influence from contact with a hydrocoel. Otherwise, in both species, V2 formed a skeletal element that has not been previously described in this location. Because of the location, shape, and timing of its formation, V2, like V1, appears to have the innate ability of the larval ectoderm to contribute to juvenile rudiment formation.     

Behavioral ecology of Strongylocentrotus droebachiensis (Muller) (Echinodermata: Echinoidea)

Summary Field observation indicated that extensive aggregation behavior occurs in Strongylocentrotus droebachiensis, an important echinoid herbivore in rocky subtidal zones of northern New England. The relationship of this behavior to the behavioral ecology of this species was investigated. Laboratory studies indicated that urchins are chemically sensitive to the stimuli of food and other urchins. Quantitative field experiments with tagged animals and transects showed the existence of two types of aggregations: feeding and nonfeeding, which appear to be pervasive phenomena in the biology of S. droebachiensis.It is concluded that, depending upon their hunger state, urchins shuttle between feeding aggregations in exposed locations and non-feeding aggregations located in relatively sheltered areas (where they spend most of their time). Individuals of S. droebachiensis aggregate actively through response to other urchins in the form of chemotaxis. The aggregative behavior may be maintained for its selective value in terms of defense from predation and, to lesser extents, facilitation of feeding and breeding.     

Gonadal growth and reproduction in the commercial sea urchin Paracentrotus lividus (Lamarck, 1816) (Echinodermata: Echinoidea) from southern Spain

The reproductive state of six populations of the commercial sea urchin P. lividus was investigated. Specimens were sampled monthly from six sites of southern Spain. The development of the gonad during the period of study was measured using histological methods and categorised by four stages for females and two for male specimens. A Gonadosomatic Index was also established. The six populations exhibited similar reproductive patterns, having mature gonads during almost all months while reconstitution and spent were present mainly in autumn. The population from Sancti Petri had the highest Gonadosomatic Index value. Higher food availability may explain this difference.     

Polykristalliner Calcit bei Seeigeln (Echinodermata,Echinoidea)

Zusammenfassung Es wurde erstmals für Echinodermen primär polykristalliner Calcit nachgewiesen, und zwar im Cortex der Primärstacheln der Cidaridae, dem sekundären Zahnskelet von Clypeaster und in den akzessorischen Kalkstrukturen, die im Kauabschnitt die Furche der Diadematiden-Zähne ausfüllen. Es gibt bei anderen Seeigelfamilien keine Bildungen, die dem Cidariden-Cortex oder den akzessorischen Kalkstrukturen der Diadematiden homolog sind. Das polykristalline sekundäre Zahnskelet von Clypeaster ist dagegen dem monokristallinen sekundären Zahnskelet anderer Seeigel homolog.Der Mg-Gehalt des Calcits liegt in den feinkristallinen Zonen (mit Ausnahme des Cortex) im allgemeinen höher; die höchsten Werte finden sich in den Steinteilen der Zähne, gleichgültig ob das sekundäre Zahnskelet mono- oder polykristallin ist.Polykristalline Teile sind im allgemeinen härter als monokristalline Teile. Die Steinteile der Seeigelzähne sind die härtesten Skeletteile von Echinodermen überhaupt; ihre VickersHärte übertrifft weit diejenige von solidem Calcit. Im Steinteil ist das feinkristalline Gefüge von Calcit eng mit organischer Matrix verbunden, und es wird vermutet, daß darauf die besonders hohe Härte der Steinteile beruht.

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Polycristalline calcite in sea urchins

Summary For the first time primary polycrystalline calcite in Echinoderms is shown in the cortex of primary spines of Cidaridae, in the secondary tooth skeleton of Clypeaster and in the accessory calcareous structures filling the crevice fold in the chewing areas of Diadematoidae teeth. Other Echinoid families lack formations homologous to the cortex of Cidaridae and accessory calcareous structures of Diadematoidae. On the other hand the polycrystalline secondary tooth skeleton of Clypeaster is homologous to the monocrystalline one of the other sea urchins.With the exception of cortex the Mg-content in calcite—analyzed by microprobe and X-ray powder method—is generally greater in macrocrystalline parts. The highest Mg-contents are found in the stone parts of teeth irrespective of whether the secondary tooth skeleton is monocrystalline or polycrystalline.Polycrystalline parts are usually harder than monocrystalline ones. The stone parts of Echinoid teeth are the hardest skeleton parts of Echinoderms on the whole; their hardness is much greater than that of solid calcite. It is supposed that the strong interlacing of the microcrystalline calcite and organic matter causes the enormous hardness of the stone part.

Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Abnormal growth patterns in the sea urchin Tripneustes CF. Gratilla (L.) under pollution (Echinodermata,Echinoidea)

Two populations of deformed sea-urchins were found in the vicinity of a combined power and desalination plant in the Gulf of Eilat (Aqaba), Red Sea. This area is highly polluted by thermohaline and heavy metal ion effluents. More than 60% of the urchins showed irregular bulging of the aboral half of the test. The height (H) to diameter (D) ratio for the most affected population was 0.70 (±0.15) compared with 0.53 (±0.04) for a normal population, away from the powerplant. In extremely deformed individuals the H/D ratio reached 1.4. The deformed sea-urchins had a wider peristome aperture and larger Aristotle's lantern, and fewer interambulacral plates than a normal urchin of the same diameter.Growth rate of highly deformed urchins in aquaria, under non-polluted conditions, was very low. In 2 months their average diameter increased by only 0.2 mm, compared with 7–8 mm in normal or slightly deformed urchins, under the same conditions.Growth lines in deformed urchins's plates indicate excessive growth in the meridional direction which is normally much less. It is suggested that controlled growth was disrupted. Some possible controlling mechanisms are discussed.     

Haltung und Zucht der Geißelspinne (Damon variegatus) (Phrynichidae) (Chelicerata: Amblypygi)

In the temporary carboniferous hall of „GONDWANA - Das Prähistorium“ in Germany, whip spiders (Damon variegatus) were kept and bred. 3,3 were purchased from a dealer in summer 2007. The night active spiders were kept together in a terrarium. No aggresions were observed. The animals were fed little locusts, young hissing cockroaches, and crickets. In 2008 courtship display and mating could be observed. The biggest male was the only active one. After 7 days after the intake of the spermatophores all females carried eggsacs under the ophistosoma. Each egg sack contained approximately 25-30 eggs. During the carrying phase only one female feasted on a cricket. 98 days after the egg sack was produced the young spiderlings hatched. The coloration of the spiderlings was completely different from the coloration of the adults. The spiderlings were carried by the female for the next 9 days. Then, the spiderlings moulted and left the female immediately. The adults were now separated from their young. The three groups of spiderlings stayed together in a smaller terrarium. However, each group from each female stayed separate from the others for weeks. The second moulting took place 41 days after the first one. No cannibalism was observed. Obviously Damon variegatus seems to be a spider with a social touch. If the terrarium and the climate conditions inside are adapted to the needs of the whip spiders keeping and breeding is possible. The species is highly attractive to visitors.     

Histology of the globiferous pedicellariae of Psammechinus miliaris (Echinodermata: Echinoidea)

The general morphology, function and histology of the globiferous pedicallariae of the regular sea-urchin, Psammechinus miliaris are described. There is a ganglion under the sensory epidermis on the inner surface of each jaw. This is the first discovery of a ganglion in any echinoid.
The regeneration of globiferous pedicellariae in laboratory conditions takes 25 to 40 days.     

Pourtalesiid sea urchins (Echinodermata: Echinoidea) of the northern Mid-Atlantic Ridge

Five species of deep-sea pourtalesiid echinoids were collected by the RV G.O. Sars MAR-ECO expedition to the northern part of the Mid-Atlantic Ridge: Solenocystis imitans new genus, new species, Echinosigra phiale (Thomson, 1872), Echinosigra (Echinogutta) fabrefacta Mironov, 1974, and two unidentified species of Pourtalesia. Three types of plastron plating are distinguished within the family Pourtalesiidae; in Solenocystis new genus the plating is similar to that in Spatagocystis, a genus known only from the deep-sea Antarctic. Each of three groups with distinctive plastron plating shows a continuous morphological gradation from a less elongated test without subanal rostrum to an extraordinarily elongated test with large rostrum and wide subanal fasciole; the ratio width/length of test varies from 0.86 to 0.12. Some morphological characters of the pourtalesiids are regarded as adaptations to burrowing and appear to be correlated with test elongation. The family is represented in the northern Atlantic only by 'advanced' genera (Echinosigra, Pourtalesia and Solenocystis) with very elongated tests.     

Phylogeny of Early Cretaceous spatangoids (Echinodermata: Echinoidea) and taxonomic implications

A phylogenetic analysis of 36 species provides a test for the taxonomy and the history of Early Cretaceous spatangoids. Most taxonomic units from genera to suborders are consistent with the proposed phylogenetic framework. We retain Hemiasterina, Micrasterina, Hemiasteridae, Schizasteridae, Hemiaster , Heteraster , Mecaster , and Periaster as original monophyletic groups. However, all of these clades originate without the classical apomorphies normally ascribed to them. We suggest a revision of their diagnoses and of the generic attributions of basal species. Some ill-defined, 'primitive', and paraphyletic taxa are recognised: Toxaster , Epiaster , Palhemiaster , and Toxasteridae. Even if they do not have phylogenetic meaning, they are retained here, pending a more complete revision.     

Thyroid hormones determine developmental mode in sand dollars (Echinodermata: Echinoidea)

Evolutionary transitions in larval nutritional mode have occurred on numerous occasions independently in many marine invertebrate phyla. Although the evolutionary transition from feeding to nonfeeding development has received considerable attention through both experimental and theoretical studies, mechanisms underlying the change in life history remain poorly understood. Facultative feeding larvae (larvae that can feed but will complete metamorphosis without food) presumably represent an intermediate developmental mode between obligate feeding and nonfeeding. Here we show that an obligatorily feeding larva can be transformed into a facultative feeding larva when exposed to the thyroid hormone thyroxine. We report that larvae of the subtropical sand dollar Leodia sexiesperforata (Echinodermata: Echinoidea) completed metamorphosis without exogenous food when treated with thyroxine, whereas the starved controls (no thyroxine added) did not. Leodia sexiesperforata juveniles from the thyroxine treatment were viable after metamorphosis but were significantly smaller and contained less energy than sibling juveniles reared with exogenous food. In a second starvation experiment, using an L. sexiesperforata female whose eggs were substantially larger than in the first experiment (202+/-5 vs. 187+/-5 microm), a small percentage of starved L. sexiesperforata larvae completed metamorphosis in the absence of food. Still, thyroxine-treated larvae in this experiment completed metamorphosis faster and in much higher numbers than in the starved controls. Furthermore, starved larvae of the sand dollar Mellita tenuis, which developed from much smaller eggs (100+/-2 microm), did not complete metamorphosis either with or without excess thyroxine. Based on these data, and from recent experiments with other echinoids, we hypothesize that thyroxine plays a major role in echinoderm metamorphosis and the evolution of life history transitions in this group. We discuss our results in the context of current life history models for marine invertebrates, emphasizing the role of egg size, juvenile size, and endogenous hormone production for the evolution of nonfeeding larval development.     

Phylogeny of Cidaroida (Echinodermata: Echinoidea) based on mitochondrial and nuclear markers

We present the first molecular phylogeny of Cidaroida, one of the most problematic groups within the echinoids. Two genes??the nuclear ribosomal gene 28?S rRNA and the mitochondrial protein-encoding gene COI??were obtained from 21 specimens representing 17 genera and 20 species, among which 13 species belong to Cidaroida. Phylogenetic analyses of the combined molecular data using parsimony and maximum likelihood optimality criteria resulted in a well-resolved phylogeny. Our results are broadly compatible (with the notable exception of Cidaris cidaris) with previous results obtained from morphological data. We find that Cidaroida represent a monophyletic group sister to the non-cidaroid Echinoidea. The family Cidaridae sensu Mortensen (1928) and Fell (1966) is paraphyletic because of the placement of Psychocidaris ohshimai as sister-group to Histocidaris elegans. Inside the Stylocidarina, we show that the two Atlantic species Stylocidaris affinis and Stylocidaris lineata constitute a well-supported clade. However, these two taxa could also represent two morphotypes within a single species showing high morphological variation.     

Abundance of Diadema antillarum (Echinodermata: Echinoidea) in the coasts of Venezuela

Diadema antillarum is a shallow-water sea-urchin from the tropical Atlantic whose populations almost disappeared in 1983-84 because of widespread mortalities which reached 87-100 %. In Venezuela, urchin population densities before the mortality event were comparable to those of other Caribbean regions; however, later abundancies remain unknown. The objectives of this study were to evaluate the recent densities of certain D. antillarum populations along the Venezuelan coasts and compare the densities at the Parque Nacional Mochima before and after the mortality. At each location urchin densities were determined by means of transects using 1 m2-quadrats as sampling units. The highest mean densities were observed at the sites on the central coast: Ensenada de Oricao, 0.28 ind/m2 (2002) and 1.05 ind/m2 (2003), and Chichiriviche de la Costa, 0.84 ind/m2 (2002) and 0.74 ind/m2 (2003). In Mochima, the mean density before the mortality for D. antillarum oscillated between 0.28 and 4 ind/m2, after the mortality event the mean density varied between 0.15 ind/m2 (2000) and 0.47 ind/m2 (2000). The populations of D. antillarum studied at Parque Nacional Morrocoy and Refugio de Fauna Silvestre Cuare showed highest densities at Playuela (0.43 ind/m2) and Cayo Sur (0.95 ind/m2) respectively, whereas other sites showed densities below 0.1 ind/m2. The density registered at Playuela in 2003 is lower than that reported before the mortality event (0.58-3.64 ind/m2). The density for Parque Nacional Archipiélago de Los Roques, specifically for the Arrecife de Herradura remained constant between 2002 and 2003 with values between 0.22-0.23 ind/m2 respectively. To conclude, the sea urchin abundancies observed at most of the Venezuelan coastal sites that we studied were higher than those reported for other areas of the northern Caribbean, even though the values have not yet returned to those preceding the 1984 mass-mortality event, due to the slow recovery of the populations.     

The effect of temperature on feeding and growth characteristics of the sea urchin Lytechinus variegatus fed a formulated feed

We investigated the effect of culture temperature on feed intake, absorption, organismal growth, and tissue production of Lytechinus variegatus by culturing individuals at three different temperatures representing the normal range of temperature exposure in wild populations in the northern Gulf of Mexico. Large L. variegatus (ca. 42 + 0.6 mm diameter, 36 + 1.3 g wet weight, n = 97) were collected at St. Joseph Bay, Florida, in October 2001. Eight sea urchins were held individually in 1-L containers within an 80-L aquarium with recirculated synthetic seawater at 32-ppt salinity. Three aquaria with the containers were each placed in three incubators at temperatures of 16, 22, or 28 °C for 8 weeks. Sea urchins held at 22 °C had the highest rate of feed intake. Feed intake in individuals held at 16 °C decreased significantly during the first 2 weeks of exposure and then increased to values not significantly different from those held at 28 °C by week 6. The dry matter absorption efficiency of individuals held at 28 °C was significantly higher than those held at 16 °C or 22 °C. The percentage of organic matter in the feces did not vary significantly with temperature. Individuals increased significantly in diameter, wet weight, and gonad weight at all temperatures. The wet weights of individuals held at 22 °C were significantly higher than those held at 28 °C or 16 °C, which did not differ significantly. The gut weight varied inversely with temperature. The wet weight of gonads of individuals held at 22 °C was significantly higher than those held at 28 °C, but neither differed significantly from those held at 16 °C. Production efficiencies, both organismal and gonadal, were inversely proportional to temperature, indicating that the overall metabolic cost of production increased with increasing temperatures. Organism production efficiencies were lower and gonad production efficiencies were higher than those found in small sea urchins, emphasizing that patterns of nutrient allocation vary between small and large sea urchins. Physiological processes associated with feed intake, absorption, and nutrient allocations vary with temperature, but allow the sea urchins to maintain growth and gonad production at a variety of temperatures. These data suggest that temperatures near the upper limits do not promote efficient use of resources, an important consideration for future commercial culture. Since gonad (roe) production is the ultimate goal of many aquaculture operations, gonad production efficiencies will provide a valuable tool for evaluating the efficacy of various feeds and feeding conditions on gonad production.