Serotonergic mechanisms mediating spawning and oocyte maturation in the zebra mussel,Dreissena polymorpha

Summary

The zebra mussel, Dreissena polymorpha, is a freshwater biofouling bivalve unintentionally introduced in the 1980s into North America from Europe. Oocyte maturation (germinal vesicle breakdown, GVBD) and spawning of the zebra mussel can be triggered with serotonin (5-hydroxytryptamine, 5-HT). In pharmacological experiments to characterize the receptor mediating spawning, the serotonin receptor agonists 8-OH-DPAT, TFMPP, and 1-(1-naphthyl)piperazine were effective at stimulating spawning; whereas, 2-methylserotonin and alpha-methylserotonin had no effect. In experiments with antagonists of serotonin receptors ketanserin and propranolol had no effect; mianserin, NAN-190, and cyproheptadine had partial inhibitory effects; and methiothepin was a very effective antagonist. Metergoline had mixed agonist/antagonist properties. Ergotamine was the most effective activator of spawning in females. Compared to serotonergic receptors in other organisms, the receptors that activate spawning in zebra mussels resemble 5HTlym, 5HTdro2 and human 5HT1Dβ, which are receptors that may act both by inhibiting adenylyl cyclase and by activating phospholipase C. In zebra mussels, 5-HT and 8-OH-DPAT activate GVBD in gonad fragments, a process also initiated by manual dissection of gonad fragments. GVBD can be inhibited by pre-treatment of ovaries with forskolin and theophylline, suggesting an inhibitory role for cyclic AMP. The Ca²⁺ ionophore A23187 can trigger GVBD and polar body formation. Thus, oocyte maturation in zebra mussels may be initiated via serotonergic receptors simultaneously inhibiting adenylyl cyclase and activating Ca²⁺ mechanisms.     

The zebra mussel (Dreissena polymorpha), a new pest in North America: reproductive mechanisms as possible targets of control strategies

Summary

The zebra mussel (Dreissena polymorpha) has spread rapidly in temperate fresh waters of North America since its introduction into the Great Lakes in 1985 or 1986. It attaches to hard substrates, forming layers, occluding water intakes, encrusting and killing native mussels, filtering algae in competition with other planktivores, and possibly interfering with fish spawning. It reproduces prolifically, suggesting that an approach to its control may be by controlling its reproduction. Previous literature suggests that spawning in bivalves is regulated by both environmental and internal chemical cues. A suggested sequence is that phytoplankton chemicals initially trigger spawning; chemicals associated with gametes provide a species-specific pheromonal positive feedback for spawning; and the response to environmental chemicals is mediated internally by serotonin (5-HT). The role of 5-HT in zebra mussels is under investigation. Both males and females can be induced to spawn by either injection or external application of 5-HT. The response can also be activated by 8-hydroxy-2-(di-n-propylamino)-tetralin, an agonist at 5-HT_1A receptors. HPLC analysis has detected 5-HT as the major biogenic amine in both male and female gonads. 5-HT immunocytochemistry demonstrates nerves containing serotonergic fibers innervating gonads of both males and females, with prominent varicosities surrounding the follicles in both sexes. A role of 5-HT in mediating spawning responses in zebra mussels is thus strongly supported. These studies have shown that reproductive behavior of zebra mussels can be modified by outside chemicals, a property that may be exploited for purposes of control.     

Variations in the Reproductive Cycle of Dreissena Polymorpha in Europe, Russia, and North America

SYNOPSIS. The reproductive cycle of the zebra mussel {Dreissenapolymorpha) is highly variable throughout its range in Europe,Russia², and North America. The environmental factors influencingthis variation are poorly understood, but successful reproductionis occurring in areas where it was initially believed that adultzebra mussels could not survive (i.e., southern United States).The differences in mussel reproduction occurring from site-to-sitemake it difficult to predict timing of specific events, suchas the start of larval production, that are important in initiatingcontainment or control procedures. For example, the amount oftime required for a fertilized egg to develop into a juvenilemussel can be as short as 8 days, or as long as 240 days. Releaseof gametes by adults can be a highly synchronized event, focusedover a 1–2 week period, or it can be completely non-synchronized,occurring throughout the year. Zebra mussels in some localitiesstart spawning at water temperatures of 12–13°C, butdo not start until water temperatures reaches 22°C at othersites. While some of this variability in reproductive behaviorstems from mussel adaptation to local conditions, part is dueto difficulties in sampling these events. It is difficult todetermine reproductive success of a specific population becauseof the problems in separating locally produced larvae from larvaedrifting in from other areas. Further research is needed notonly on the relationship between reproduction and environmentat the community level, but also on the variability in responseof individual mussels.     

Activation of spawning in zebra mussels by algae-, cryptomonad-, and gamete-associated factors

In zebra mussels (Dreissena polymorpha) simultaneous release of gametes and peaks in larval densities at particular locations suggest that spawning is triggered by synchronizing stimuli. Furthermore, spawning tends to occur only after an adequate environmental temperature is reached. To test the hypothesis that phytoplankton and gamete-associated chemicals initiate spawning in zebra mussels and that the responsiveness to such chemicals is affected by ambient temperature, the spawning response of zebra mussels to extracts from algae, a cryptomonad, and a cyanobacterium and to water associated with released gametes was assayed in animals acclimated to 12 ^C and 17 ^C. For animals held at 12 ^C, only serotonin, a known activator of bivalve spawning used as a positive control, stimulated spawning. However, for animals acclimated to 17 ^C, extracts made from a diatom (Phaeodactylum), a brown alga (Fucus), and a cryptomonad (Rhodomonas) stimulated spawning in both sexes; extracts from green algae (Platymonas and Dunaliella) and a cyanobacterium (Oscillatoria) did not cause spawning. Water associated with either released sperm or eggs elicited spawning in both females and males. Positive controls, stimulated with serotonin, spawned at a high (>90%) rate, whereas no negative control spawned. Thus, phytoplankton chemicals and gamete-associated factors may have a role in synchronizing spawning in zebra mussels once adequate ambient temperature is reached.     

Putative serotonin reuptake inhibitor-induced spawning and parturition in freshwater bivalves is inhibited by mammalian 5-HT2 receptor antagonists

Selective serotonin reuptake inhibitors (SSRIs) can mimic the physiological actions of serotonin, and in bivalve molluscs they induce zebra mussel spawning and fingernail clam parturition. We have elucidated further the pharmacology of SSRI-induced spawning and part-urition by blocking these reproductive processes with two mammalian 5-HT(2) receptor antagonists, cyproheptadine and mianserin. These two antagonists were potent inhibitors of both spawning and parturition induced by the SSRIs fluvoxamine, fluoxetine, and zimelidine. In zebra mussels, both cyproheptadine and mianserin significantly blocked spawning induced by fluvoxamine and by zimelidine. In the fingernail clams Sphaerium spp., both cyproheptadine and mianserin blocked fluvoxamine-induced parturition. A possible mechanism of action for SSRI-induced spawning and parturition in bivalves is suggested.     

ENDOSYMBIONTS OF DREISSENA POLYMORPHA IN IRELAND: EVIDENCE FOR THE INTRODUCTION OF ADULT MUSSELS

Although zebra mussels (Dreissena polymorpha) have invaded watersacross Europe for over 200 years, they colonized Ireland onlywithin the past decade. To test the hypothesis that Irelandwas colonized by adult D. polymorpha, we examined mussels fromdifferent sites along the Lower Shannon River system in Irelandfor the presence of host specific and generalist endosymbionts.Withinthe mantle cavity and/or associated with zebra mussel tissueswe found species specific-ciliates (Conchophthirus acuminatusand Ophryoglena hemophaga) and generalist symbionts (the ciliateAncistrumina limnica, nematodes, oligochaetes and chironomids).We found a significant difference in the prevalence of symbiontsamong sites, but all mussels at all sites harboured one specialistspecies C. acuminatus, and all of the mussels at three of thefour sites also had the second specialist, O. hemophaga. Thus,with the introduction of D. polymorpha into Ireland, at leasttwo additional species, their host-specific symbionts C. acuminatusand O. hemophaga, have also been introduced. The presence ofthese symbionts in Ireland supports the hypothesis that adultzebra mussels were introduced into Ireland, rather than larvalstages. This contrasts with the introduction of zebra musselsto North America, where adult zebra mussels are devoid of host-specificsymbionts. (Received 8 June 2005; accepted 7 November 2005)     

Grazing on colonial and filamentous, toxic and non-toxic cyanobacteria by the zebra mussel Dreissena polymorpha

Colony forming and toxic cyanobacteria form a problem in surfacewaters of shallow lakes, both for recreation and wildlife. Zebramussels, Dreissena polymorpha, have been employed to help torestore shallow lakes in the Netherlands, dominated by cyanobacteria,to their former clear state. Zebra mussels have been presentin these lakes since they were created in the 19th century bythe excavation of peat and are usually not considered to bean invasive species. Most grazing experiments using Dreissenahave been performed with uni-cellular phytoplankton laboratorystrains and information on grazing of larger phytoplankton taxahardly exists. To gain more insight in to whether D. polymorphais indeed able to decrease cyanobacteria in the phytoplankton,we therefore performed grazing experiments with zebra musselsand two species of cyanobacteria, that greatly differ in shape:colony forming strains of Microcystis aeruginosa and the filamentousspecies Planktothrix agardhii. For both species a toxic anda non-toxic strain was selected. We found that zebra musselscleared toxic Planktothrix at a higher rate than non-toxic Planktothrix,toxic or non-toxic Microcystis. Clearance rates between theother strains were not significantly different. Both phytoplanktonspecies, regardless of toxicity, size and shape, were foundin equal amounts (based on chlorophyll concentrations) in theexcreted products of the mussels (pseudofaeces). The resultsshow that zebra mussels are capable of removing colonial andfilamentous cyanobacteria from the water, regardless of whetherthe cyanobacteria are toxic or not. This implies that the musselsmay be used as a biofilter for the removal of harmful cyanobacterialblooms in shallow (Dutch) lakes where the mussels are alreadypresent and not a nuisance. Providing more suitable substratefor zebra mussel attachment may lead to appropriate mussel densitiescapable of filtering large quantities of cyanobacteria.     

Seasonal reproductive cycle in the mussel,Lamellidens corrianus

Seasonal reproductive cycle of the freshwater mussel, Lamellidens corrianus has been studied. These mussels are functional or simultaneous hermaphrodites. The spawning was at its peak during the months of September to December. The gonads were in growing stages with reduced gonadal activity during January to April, whereas the maturation of gonads was found to be intense during May to August.     

Increased gonad ratio in the blue mussel, Mytilus edulis, exposed to starfish predators

The presence of invertebrate predators can induce morphological, physiological, and behavioural changes in sessile marine prey species. In this study, I investigated whether the risk of predation can alter the reproductive effort in the blue mussel Mytilus edulis. In the field, I cultured blue mussels in the presence and in the absence of the starfish, Asterias rubens, using enclosures with two compartments. Two experiments were performed with field periods of 19 and 30 days, respectively. After the field exposure, the mussels were examined for weight of flesh and gonad tissue. In both experiments, the starfish-exposed mussels developed a significantly larger gonad to flesh ratio, compared with that of the unexposed control mussels. After 30 days, the exposed mussels had also lost flesh weight significantly, probably due to spawning. These changes suggest that M. edulis accelerates the gonad development and/or increases its reproductive effort in the presence of predators. Predator-inducible adjustment of life-history characters should be a beneficial trait for this sessile species, that typically encounters a high and variable predation pressure. This revised version was published online in August 2006 with corrections to the Cover Date.     

Zebra mussel adhesion: structure of the byssal adhesive apparatus in the freshwater mussel, Dreissena polymorpha

The freshwater zebra mussel (Dreissena polymorpha) owes a large part of its success as an invasive species to its ability to attach to a wide variety of substrates. As in marine mussels, this attachment is achieved by a proteinaceous byssus, a series of threads joined at a stem that connect the mussel to adhesive plaques secreted onto the substrate. Although the zebra mussel byssus is superficially similar to marine mussels, significant structural and compositional differences suggest that further investigation of the adhesion mechanisms in this freshwater species is warranted. Here we present an ultrastructural examination of the zebra mussel byssus, with emphasis on interfaces that are critical to its adhesive function. By examining the attached plaques, we show that adhesion is mediated by a uniform electron dense layer on the underside of the plaque. This layer is only 10-20 nm thick and makes direct and continuous contact with the substrate. The plaque itself is fibrous, and curiously can exhibit either a dense or porous morphology. In zebra mussels, a graded interface between the animal and the substrate mussels is achieved by interdigitation of uniform threads with the stem, in contrast to marine mussels, where the threads themselves are non-uniform. Our observations of several novel aspects of zebra mussel byssal ultrastructure may have important implications not only for preventing biofouling by the zebra mussel, but for the development of new bioadhesives as well.     

Aspects of the reproductive biology of the South African polychaete,Arenicola loveni loveni (Kinberg 1866)

Summary

The annual cycles of gametogenesis, spawning and fertilization biology were investigated in the South African polychaete Arenicola loveni loveni (Kinberg, 1866). Sexes are strictly separate (gonochoric) with a 1:1 ratio and gametogenesis takes place in the coelomic cavity. Reproduction is iteroparous and occurs during a synchronized annual epidemic spawning event during the summer months of February or March, which lasts no longer than 2 weeks at around the time of highest annual seawater temperatures. A slight inter-annual variation in spawning time was observed between 2003 and 2004. Annual fecundity was estimated as 311,200 (± 82,040 standard deviations) oocytes per female per year. Oocytes are arrested at prophase I of the first meiotic division and require a maturation step prior to spawning and subsequent fertilization. Mean spawned oocyte diameter was 141.16 μm ± 0.43 SE. Sperm develop as morulae in the coelomic cavity and also require a maturation step prior to spawning. Spawning can be achieved by injection of prostomial homogenate in both males and females and by injection with 8,11,14- e cosatrienoic acid for males and can be delayed by temperature manipulation. Oocyte maturation and sperm maturation can be induced in vitro by incubating in prostomial homogenate.     

LIGHT, GRAVITY AND CONSPECIFICS AS CUES TO SITE SELECTION AND ATTACHMENT BEHAVIOUR OF JUVENILE AND ADULT DREISSENA POLYMORPHA PALLAS, 1771

Dreissena polymorpha, the zebra mussel, is one of the mostimportant components of the aquatic ecosystems it inhabits,due to high densities and filtration rates of this fouling organism.This laboratory study deals with several physical and biologicalfactors influencing zebra mussel juveniles (<10 mm) and adults(>10 mm) site selection behaviour and their byssal attachmentto substrate. Mussels preferred black substrate rather thanwhite. This preference was found to be stronger in smaller individuals.Furthermore, all mussels, independently of their size, selectedshadowed sites and avoided illuminated ones. Large mussels attachedmore often in darkness than in light, while for small specimenslight conditions made no difference. Large mussels tended toattach to lower parts of vertical test-tubes, while the numbersof small specimens in both halves of the test-tubes were similar.If initial conditions, used by mussels to select their attachmentsite, were changed by turning the test-tubes upside down, smallindividuals detached and looked for a new site more frequentlythan large ones. Large mussels attached more often in higherdensities of conspecifics, but physical contacts among conspecificsand their size composition seemed to have no impact on attachment.This study shows that many environmental stimuli influence locomotionand attachment of post-metamorphic zebra mussels. Therefore,the behaviour of this group may impact upon adult distributionobserved in field. Preferences exhibited by mussels lead tothe creation of dense aggregations in dark, deep places, providing protectionagainst dessication, predators and excessive water flow. (Received 18 April 2000; accepted 5 October 2000)     

Effects of temperature and temperature acclimation on survival of zebra mussels (Dreissena polymorpha) and Asian clams (Corbicula fluminea) under extreme hypoxia

Following acclimation to 5°, 15° or 25°C for 14days, samples of 30 Dreissena polymorpha (zebra mussels) andCorbicula fluminea (Asian clams) were held in either aerated(control) or extremely hypoxic N₂ gassed water (PO2 < 3%of full air saturation). Mortality was negligible in all aeratedcontrols. Mean hypoxia tolerance in D. polymorpha ranged from3–4 days at 25°C to 38–42 days at 5°C. Hypoxiatolerance time of zebra mussels increased significantly withdeclining test temperature (P < 0.001) and increasing acclimationtemperature (P < 0.001). Larger zebra mussels were more tolerantthan smaller individuals. Asian clams were 2–7 times moretolerant of hypoxia than zebra mussels, surviving a mean of11.8 and 35.1 days at 25°C and 15°C, respectively, andwithout mortality for 84 days at 5°C, and were not influencedby temperature acclimation. At 25°C, larger specimens ofAsian clams were less tolerant of hypoxia than smaller individuals.Both species are amongst the least hypoxia tolerant freshwaterbivalve molluscs, reflecting their prevalence in well-oxygenatedshallow water habitats. Prolonged exposure to extreme hypoxiamay provide an efficacious control strategy, particularly forD. polymorpha (Received 12 January 1998; accepted 30 September 1998)     

Growth and reproductive cycle of the zebra mussel in the River Rhine as studied in a river bypass

In a Rhine River bypass built at a water pollution control station, the growth and reproductive cycle of the zebra mussel (Dreissena polymorpha) were studied over a period of nearly 3 years. The experimental setup offered the possibility to manipulate the temperature of the continuously passing river water and to calculate influences of temperature and food on the growth of individually marked mussels. Shell growth rates were a function of algal density (measured as chlorophyll a). This function followed saturation curves with temperature-dependent plateau levels, and saturation occurred at values above 40 μg/l chlorophyll a. The relationship between shell growth rate and temperature followed the courses of bell-shaped curves with algal-food-dependent heights. No clear correlations could be found between shell growth and other environmental factors such as dissolved organic matter, total organic content, seston, and turbidity. Two spawning periods per year characterized the reproductive cycle of the zebra mussels at the study site. The first one covered nearly 3 months (April to July) comprising several spawning events. The second shorter one occurred in August. Young mussels of the first spawning period settled at the end of May, and they were able to reproduce a few months later (end of August) when their shell length exceeded 9 mm. This first spawning period of the young-of-the-years coincided with the second spawning period of the older generations. Through the combined analysis of the results of both body weight determinations and shell growth experiments, it was possible to model mussel growth over a period of nearly 3 successive years, and to suggest the presence of an endogenous factor that might be involved in determining the start of the annual shell growth period. Received: 8 April 1997 / Accepted: 3 November 1997     

Plasticity of fertilization rates under varying temperature in the broadcast spawning mussel,Mytilus galloprovincialis

Oceans are a huge sink for the increased heat associated with anthropogenic climate change, and it is vital to understand the heat tolerance of marine organisms at all life stages to accurately predict species’ responses. In broadcast spawning marine invertebrates, reproduction is a vulnerable process in which sperm and eggs are released directly into the open water. Gametes are then exposed to fluctuating environmental conditions that may impact their fertilizing capacity. Using the broadcast spawning Mediterranean mussel, Mytilus galloprovincialis, as a model species, we performed blocks of factorial mating crosses to assess the variance in fertilization rates among individuals under both ambient and elevated temperatures. Overall, we found a small, but significant decline in fertilization rates with elevated temperatures. However, there was substantial plasticity in responses, with particular mussels having increased fertilization under elevated temperatures, although the majority showed decreased fertilization rates. Our results suggest possible future reproductive costs to ocean warming in M. galloprovincialis, although it is also possible that genetic variation for thermal sensitivity may allow for adaptation to changing environmental conditions.     

A Review of the Biology and Ecology of the Quagga Mussel (Dreissena bugensis), a Second Species of Freshwater Dreissenid Introduced to North America

SYNOPSIS. North America's Great Lakes have recently been invadedby two genetically and morphologically distinct species of Dreissena.The zebra mussel (Dreissena polymorpha) became established inLake St. Clair of the Laurentian Great Lakes in 1986 and spreadthroughout eastern North America. The second dreissenid, termedthe quagga mussel, has been identified as Dreissena bugensisAndrusov, 1897. The quagga occurs in the Dnieper River drainageof Ukraine and now in the lower Great Lakes of North America.In the Dnieper River, populations of D. polymorpha have beenlargely replaced by D. bugensis; anecdotal evidence indicatesthat similar trends may be occurring in the lower LaurentianGreat Lakes. Dreissena bugensis occurs as deep as 130 m in theGreat Lakes, but in Ukraine is known from only 0–28 m.Dreissena bugensis is more abundant than D. polymorpha in deeperwaters in Dneiper River reservoirs. The conclusion that NorthAmerican quagga mussels have a lower thermal maximum than zebramussels is not supported by observations made of populationsin Ukraine. In the Dnieper River drainage, quagga mussels areless tolerant of salinity than zebra mussels, yet both dreissenidshave acclimated to salinities higher than North American populations;eventual colonization into estuarine and coastal areas of NorthAmerica cannot be ignored.     

Sexual maturity in young Dreissena polymorpha from Lake Como (N. Italy)

Summary

The reproductive biology of the zebra mussel Dreissena polymorpha was studied during 1999 in a natural population living in Lake Como (N. Italy), with particular reference to the differences in behavior between young and adult mussels. Histological analysis of gonads was used to study the gametogenic cycles and to evaluate when the young reached sexual maturity. Signs of active gametogenesis were detected in ≥4 mm shell length mussels. Mature gametes and the first spawning events were observed in ≥5 mm males and in ≥6 mm females. The gonad feature of ≥7 mm mussels was similar to that of the adults. Size frequency distributions revealed that the new generation (0⁺) first spawned in the year following birth, but only after winter. The length of larval time seemed to vary greatly depending on the spawning period, in addition to the environmental conditions. The first settled mussels ≥500 μm in shell length were observed 3–4 months after the first depositions. Water temperature seemed to be the most important factor which regulated sexual phenomena of both young and adult mussels. Its role in the onset of the spawning events was confirmed, but it is important to point out that temperature is not enough to trigger reproduction since gamete release is impossible without sufficient energy, especially for young mussels.     

Maturation, Spawning and Development in Siphonaria Pectinata Linnaeus (Gastropoda: Pulmonata) at Gibraltar

A comparative study of some aspects of the reproductive biologyof Siphonaria pectinata was carried out at Gibraltar. Maturationoccurred at 5-6 mm shell length corresponding to an age of 6-12months. At the more sheltered Atlantic site spawning was evidentfrom March to June 1992 with peak spawning occurring in May-June1992. The patterns of spawning activity for the following twoyears were very similar. Egg ribbons were very scarce at themore exposed Mediterranean site and were found in considerablyless quantities than at the Atlantic site. Individuals at bothsites contained mature eggs in their gonads at the onset, andthen throughout the duration of the spawning season, but exhibitedspent gonads by October. The spawn of S. pectinata consistedof small oval ribbons filled with numerous egg-shaped capsulesspaced irregularly in a jelly-like matrix. Spawn was depositedmainly away from the home scar in shallow depressions, cracksand crevices and between barnacles and mussels. S. pectinataexhibited planktonic development. The fertilised eggs took between14 and 21 days to develop at ambient temperature (14-19°),and hatched as free-swimming planktotrophic veligers. The timingof spawning activity and the placement of egg masses in protectivemicrohabitats on the shore are discussed in terms of adaptationsto reduce desiccation stress, predation and possible food shortages.The mode of development in S. pectinata is placed in contextwith current theories regarding siphonariid ancestry and evolution. (Received 6 January 1998; accepted 25 June 1998)     

Changes in trace metal concentrations in the tissues of the White Sea mussel <Emphasis Type="Italic">Mytilus edulis</Emphasis> over the reproductive cycle

Seasonal changes in the concentrations and total amounts (contents) of Cd, Zn, and Cu in the mussel Mytilus edulis cultivated in the White Sea were studied over the reproductive cycle (prespawning, spawning, and postspawning stages). The results, when compared with published data on the closely related species M. trossulus from the Sea of Japan, suggest that the seasonal dynamics of trace element contents in mussel tissues are related to specific geochemical conditions, as well as to the dynamics of changes in soft tissue weight over the reproductive cycle. The contents of Cd and Cu in the mussels before, during, and after spawning changed similarly in the mollusks from the White Sea and the Sea of Japan. Changes in the Zn content at different stages of the reproductive cycle of the White Sea mussels were similar to those in mussels from the Sea of Japan but had smaller amplitude. The concentration of Zn in the White Sea mussels was before spawning the highest, but still lower than in mussels from the Sea of Japan. After the spawning, the Zn concentration in the White Sea mussels, in contrast to the Pacific mussels, decreased because of the redistribution of this element during the prespawning period from the somatic tissue into the gonad.     

Growth, Degrowth, and Irreversible Cell Differentiation in Aurelia aurita

Growth patterns of the Scyphomedusa Aurelia aurita from TomalesBay. California, were examined in the field and in the laboratory.Manipulation of growth patterns demonstrated that degrowth andregrowth are not constrained by initial ievelopmental stage.Although initial degrowth of certain tissues is allometric (e.g.,gonads regress in 5 to 8 days; bell diameter decreases morerapidly at first than do the oral arms), thereafter regressionappears identical to, but reversed from normal growth. Regrowthpatterns are normal. Sexual maturation in the sea does not alwaysalter subsequent capacity for degrowth or regrowth to sexualmaturity in the laboratory, because reproductive and somatictissues do not always degenerate after spawning. Gonadal tissuecan be renewed and maintained in a ripe condition in the laboratoryapparently indefinitely. Sexual maturation is a size-dependentphenomenon, not an agespecific developmental event. Spermatogenesis, once initiated, proceeds irrespective of outsideevents. Labeled spermatogonial cells can continue to differentiateto form sperm even though the gonad containing those cells,and the animal itself, show rapid degrowth. The importance ofthis decoupling of developmental events is discussed. The experimentalimportance of animals with flexible life cycles is emphasized.