Phytoplankton as a stimulus for spawning in three marine invertebrates

A distinct annual reproductive cycle with spring spawning was observed in Strongylocentrotus droebachiensis Müller, Tonicella lineata Wood and Tonicella insignis Reeve. This study gives evidence that the cue for spawning in these species is the spring phytoplankton bloom. In 1973 spawning occurred abruptly in early April at the time of the spring phytoplankton outburst, but in 1974 spawning was less abrupt corresponding to the slow development of the phytoplankton bloom in that year. Animals were collected prior to spawning and maintained in the laboratory under various temperature and light regimes: at 5 ° and 14 °C in darkness, and at 5 ° and 14 °C in light conditions similar to those in the field. These animals did not spawn when spawning occurred in the field, but animals returned to the field from the laboratory did spawn. In the laboratory a large proportion of animals spawned when they were exposed to phytoplankton collected with a 50μm mesh net. The results suggest that some substance bound to or released by phytoplankton stimulates spawning. For species with planktotrophic larvae the synchronization of spawning with the phytoplankton bloom increases the probability of favourable food and temperature conditions for the development of the larvae and juveniles.     

Communal Spawning in the Common Frog Rana temporaria– Egg Temperature and Predation Consequences

Females of the common frog, Rana temporaria, spawn synchronously at communal breeding sites, forming a communal egg mass of individual spawn clumps. A total of four spawning sites were studied. In the centre of the communal egg mass, daily maximum temperature was higher and minimum temperature lower than further out. Spawn clumps in the centre of the communal egg mass developed faster than single spawn clumps. In the centre of the egg masses, some spawn clumps suffered almost total egg mortality. This was probably the consequence of a cold spell with sub‐zero temperatures in combination with a tendency for low minimum temperatures in the centre. In a laboratory study, worm leeches, Erpobdella spp., fed readily on eggs of common frog, while horse leeches, Haemopis sanguisuga, hardly ate any eggs. At the spawning sites, most worm leeches were found in single spawn clumps, far from the communal egg mass. Females which spawned late, when most of the spawning was over, preferred to spawn in the centre of the communal egg mass. This suggests that there is a net advantage for eggs in spawn clumps laid in the centre.     

Captive rearing for Chinook salmon (Oncorhynchus tshawytscha) and Atlantic salmon (Salmo salar): the Idaho and Maine experiences

Captive rearing is a conservation strategy where juveniles are collected from the natural environment, reared to maturity in a hatchery environment, and then released back into the natural environment at maturity for volitional spawning. This strategy has been used to produce adult outplants for stock enhancement where natural escapement is poor or capture of adults is difficult. In both Idaho (Chinook salmon, Oncorhynchus tshawytscha) and Maine (Atlantic salmon, Salmo salar), captive rearing programs have been initiated as an experimental strategy to prevent cohort collapse and conserve genetic integrity of select depressed populations. In this paper, we provide an overview of these programs and describe some of the methods used to evaluate the effectiveness of this approach. Behaviors such as habitat selection, courting, and spawn timing were monitored. Data collected for both programs indicate that the captive fish display similar behaviors as their wild conspecifics in terms of habitat selection and spawning, although there were some differences in spawn timing. Evaluations of egg and fry production also indicate that captive-reared adults are successfully spawning and producing offspring. Each program is still waiting on final evaluations of reproductive success through genetic analyses of returning adults, but results so far indicate that this could be an additional captive propagation strategy for depressed populations.     

The date of spawning in populations of the Common frog (Rana temporaria) from different altitudes in northern England

A series of ponds in the Wear Valley in northern England, ranging in altitude from 46 to 838 m, were visited regularly over a four-year-period, and a record was kept of the dates on which spawn from the Common frog, Rana temporaria , was first seen in each of the ponds. On average. spawning was delayed by six days for every 100 m increase in altitude. The date of spawning at each of the ponds was fairly consistent from year to year; only in 1974 was the date of spawning noticeably different from other years. A theory was proposed which could account for the variation in the date of spawning with altitude and the annual variation in the date of spawning, observed in the present study. It is proposed that frogs from nothern England become sensitive to environmental temperature in the middle of February and emerge in the spring after experiencing a temperature-sum of 106 degree(C)-days. The onset of temperature sensitivity is probably initiated by a circannual endogenous rhythm. The time at which the rhythm switches on the temperature-sensing mechanism in frogs probably varies in different populations, and may well be locally adapted. For spawning to follow emergence, a certain minimum water temperature may be required. In the present study, most frogs in the wild spawned when the water temperature in the ponds was above 7°C, but frogs could spawn at 3·1°C. Alternative theories for the control of emergence and spawning are also discussed.     

Sound Production in Codoma ornata (Girard) (Cyprinidae)

Male Codoma ornata produce sounds associated with aggression and spawning activities during the breeding season. Females do not produce sounds. Males most often produced sounds associated with escalated displays of aggression, courtship and the spawning act. C. ornata spawn in crevices, but previously were reported to spawn as egg-clusterers in cavities. Structurally, sounds are low frequency, vary in duration according to context and are not harmonic. The mechanism of sound production is unknown.     

Facultative semelparity in capelin Mallotus villosus (Osmeridae)-an experimental test of a life history phenomenon in a sub-arctic fish

Two alternative reproductive modes are present in fishes and reflect the age-specific mortality encountered through ontogenesis. Life-history hypotheses suggest that semelparity (i.e. death after a single reproductive event) evolves when the ratio of juvenile to adult survival is relatively high. Conversely, a relatively low ratio of juvenile to adult survival will favour iteroparity (i.e. death after two or more reproductive events). Fisheries management associates capelin (Mallotus villosus) spawning with mass mortality and semelparity even though life history models developed for this species suggest that females may follow an iteroparous trajectory. Capelin may spawn either inter-tidally on the beach or offshore in deeper, ocean waters but post-spawning survival and potential iteroparity has been notoriously difficult to assess in natural populations. Through a series of aquarium experiments we tested post-spawning survivability in a beach spawning and an ocean spawning population. The findings demonstrate that capelin which spawn offshore are absolute semelparous (death of both genders) while beach spawning capelin are iteroparous irrespective of sex. Beach spawning capelin regenerated ripe gonads from one spawning season to the next and provides the first conclusive evidence that capelin is physiologically capable of an iteroparous reproductive mode. The potential physical and biological processes which generate certain reproductive patterns in capelin are summarized and discussed in relation to life history hypotheses. We suggest that capelin is a facultative semelparous species in which dynamic changes within the semelparity-iteroparity continuum may occur as a result of subtle interactions between the spawning habitat, physical forcing, and predatory pressure.     

Sperm competition affects male behaviour and sperm output in the rainbow darter

Rainbow darters, Etheostoma caeruleum, are promiscuous fish with moderate rates of group spawning (between one and five males may simultaneously mate with one female). In this study, I examined male sperm output and male willingness to spawn under different levels of sperm competition intensity. One male and one female were allowed to spawn in an aquarium where they had visual and olfactory access to one of four treatments: four males, one male, zero males, or one female. Theory predicts that males should reduce sperm output when there are more than the average number of males at a group spawning (four-male treatment) and should increase sperm output when there are fewer than average males at a group spawning (one-male treatment). Mean sperm output did not differ among treatments. However, males released more sperm when spawning in the presence of competing males (four-male and one-male treatments pooled) than when spawning in the absence of competing males (zero-male and one-female treatments pooled). Males were also most likely to forego spawning opportunities when sperm competition intensity was high. Furthermore, male willingness to spawn was size dependent. Large males were more likely to forego spawning opportunities under high sperm competition intensity. Large males may be better off waiting for future spawning opportunities when there is a lower potential for sperm competition intensity.     

A Test for Male Parental Care in a Fundulid, the Bluefin Killifish, Lucania goodei

Under field conditions, breeding male bluefin killifish, Lucania goodei, have been observed aggressively defending territories from other breeding males, non-breeding females, and minnows (mainly Notropis harperi). We performed an aquarium experiment to test whether male aggression serves to protect newly deposited eggs from predation. We allowed a male and a female to spawn in a yarn mop, removed the female, and exposed the eggs to one of four treatments (spawning male present, two minnows present, spawning mal+two minnows present, no adult fish present). Mops were censused daily for seven days. Egg predation rates were highest in the male+minnows and male only treatments. Egg predation rates in the male+minnows treatment did not differ from the predicted predation rate (sum of male only and minnows only treatments). Hence, there is no evidence for male parental care in L. goodei. In addition, we compared the egg predation rates (filial cannibalism) between males of 3 different color morphs and found no evidence for differential egg cannibalism.     

Allopaternal care in the tessellated darter,Etheostoma olmstedi (Pisces: Percidae)

Synopsis Large male tessellated darters, Etheostoma olmstedi, defend flat-bottomed rocks, the undersides of which serve as spawning substrates. Because females attach eggs directly onto bare stone, a spawning bout necessarily decreases the surface area available for further breeding at a nest. In apparent response to the female preference of ovipositing on rocks with the most bare surface, fathers abandon nests with their attached offspring to search for rocks with more uncovered surface. Egg abandonment also results after roving fathers intercept and spawn with ripe females and remain at a different rock to care for a new clutch. In essence, large males appear to maximize the number of eggs they fertilize by sequentially monopolizing breeding patches that are temporarily most desirable to females. The costs to a father of abandoning his eggs appear to be low for two reasons: (1) non-territorial males, created by a scarcity of suitable rocks, readily occupy vacant nests, and clean and aerate the abandoned eggs; and (2) after water hardening, eggs are less vulnerable to cannibalism. The maintenance of eggs by nonparental males may be both incidental to scrubbing the ceiling for their own breeding and adaptive in that viable eggs stimulate females to spawn; neither would appear to involve parental investment costs. Thus, allopaternal care in the tessellated darter may have evolved because it is performed in the selfish pursuit of spawning opportunities while entailing little, if any, of the costs normally subsumed within parental investment.     

Preliminary observations on the spawning conditions of the European amphioxus (Branchiostoma lanceolatum) in captivity

Members of the subphylum Cephalochordata, which include the genus Branchiostoma (i.e. amphioxus), represent the closest living invertebrate relatives of the vertebrates. To date, developmental studies have been carried out on three amphioxus species (the European Branchiostoma lanceolatum, the East Asian B. belcheri, and Floridian-Caribbean B. floridae). In most instances, adult animals have been collected from the field during their ripe season and allowed (or stimulated) to spawn in the laboratory. In any given year, dates of laboratory pawning have been limited by two factors. First, natural populations of these three most studied species of amphioxus are ripe, at most, for only a couple of months each year and, second, even when apparently ripe, animals spawn only at unpredictable intervals of every several days. This limited supply of living material hinders the development of amphioxus as a model system because this limitation makes it more difficult to work out protocols for new laboratory techniques. Therefore we are developing laboratory methods for increasing the number of amphioxus spawning dates per year. The present study found that a Mediterranean population of B. lanceolatum living near the Franco-Spanish border spawned naturally at the end of May and again at the end of June in 2003. Re-feeding experiments in the laboratory demonstrated that the gonads emptied at the end of May refilled with gametes by the end of June. We also found that animals with large gonads (both, obtained from the field and kept and fed at the laboratory during several weeks) could be induced to spawn in the laboratory out of phase with the field population if they were temperature shocked (spawning occurred 36 hours after a sustained increase in water temperature from 19 degrees C to 25 degrees C).     

Factors affecting the circatidal rhythm in vertical swimming of ovigerous blue crabs, Callinectes sapidus, involved in the spawning migration

Ovigerous blue crabs, Callinectes sapidus, are observed to undergo nocturnal ebb-tide transport (ETT) during their seaward spawning migration. A previous study found that females undergoing the spawning migration have a circatidal rhythm in vertical swimming, which serves as the biological basis for ETT. The present study asked three questions about this endogenous rhythm. First, does the rhythm occur in females with mature embryos regardless of whether they are undergoing ETT? Second, when exposed to a light/dark cycle in the laboratory, do ovigerous females only swim vertically at the time of ebb tide during the dark phase? Third, do attachments to the backs of ovigerous crabs affect the circatidal rhythm? The circatidal rhythm occurred in all crabs with mid-stage embryos that were prevented from undergoing ETT. The rhythm was unaffected by the light/dark cycle, which implies that migration can occur at lower light levels at depth during the day. Finally, attachments did not affect the rhythm, which suggests that tags and transmitters will not affect the spawning migration.     

Microsatellite DNA Data Indicate Distinct Native Populations of Kokanee, Oncorhynchus nerka, Persist in the Lake Sammamish Basin, Washington

Large-scale introductions of resident and anadromous salmonids from exogenous sources and urbanization have led to major changes in, and concern for the fate of, indigenous fish populations of the Lake Sammamish/Lake Washington Basin. Specifically, introductions of kokanee (the resident form of Oncorhynchus nerka) from the Lake Whatcom Hatchery and sockeye (the anadromous form of O. nerka) from Baker Lake have caused uncertainty about the ancestry of the kokanee that currently spawn in the basin. We used nine microsatellite loci to investigate the inter-relationships of kokanee populations that spawn in streams in the Sammamish sub-basin, sockeye salmon populations that share spawning areas with the kokanee, Lake Whatcom Hatchery kokanee and Baker Lake sockeye, and an outgroup, Meadow Creek kokanee, from Lake Kootenay which drains into the upper Columbia River. We observed high levels of genetic variation (5–49 alleles per locus). Explicit tests of population sub-division revealed that collections from most spawning aggregations differed from each other. Observed allele frequency distributions strongly suggest that natural spawning kokanee in the basin are not descended from recent Lake Whatcom stock introductions. We found no compelling evidence to suggest that the kokanee sampled from spawning areas within the Lake Sammamish sub-basin have resulted from, or been altered substantially by, past introductions of non-native kokanee or sockeye.     

A Sperm Spawn-Inducing Pheromone in the Silver Lip Pearl Oyster (<Emphasis Type="Italic">Pinctada maxima</Emphasis>)

Pheromones are considered to play an important role in broadcast spawning in aquatic animals, facilitating synchronous release of gametes. In oysters, the sperm has been implicated as a carrier for the spawn-inducing pheromone (SIP). In hatchery conditions, male pearl oysters (Pinctata maxima) can be stimulated to spawn through a variety of approaches (e.g. rapid temperature change), while females can only be induced to spawn through exposure to conspecific sperm, thus limiting development of targeted pairing, required for genetic research and management. The capacity for commercial production and improvement of genetic lines of pearl oysters could be greatly improved with access to a SIP. In this study, we prepared and sequenced crude and semi-purified P. maxima sperm extracts that were used in bioassays to localise the female SIP. We report that the P. maxima SIP is proteinaceous and extrinsically associated with the sperm membrane. Bioactivity from pooled RP-HPLC fractions, but not individual fractions, suggests that the SIP is multi-component. We conclude that crude sperm preparations, as described in this study, can be used as a sperm-free inducer of female P. maxima spawning, which enables for a more efficient approach to genetic breeding.     

The spawning behaviour of the endangered freshwater fish Ladigesocypris ghigii (Gianferrari, 1927)

Ladigesocypris ghigii (Gianferrari, 1927), commonly called gizani, is an endangered freshwater fish endemic to the Greek island of Rhodes. The spawning behaviour of gizani was studied in aquaria, to develop an artificial breeding technique for this endangered species. Spawning in captivity was recorded over 41 spawning days between 15 February and 14 July 2001, under constant temperature (18 or 21°C) and photoperiod 14L/10D. The number of spawning acts in a day was 82 ± 28. No aggressiveness and no territoriality were developed. The mating system of the fish was polygamous, with females spawning more than once. In captivity, eggs are laid either on aquatic plants or on gravel, while in nature fish spawn mainly on algae and plants. The latter suggests that, as far as breeding substrate is concerned, gizani is an opportunistic species using different substrates when necessary.     

Not All Offspring Are Created Equal: Variation in Larval Characteristics in a Serially Spawning Damselfish

The way organisms allocate their resources to growth and reproduction are key attributes differentiating life histories. Many organisms spawn multiple times in a breeding season, but few studies have investigated the impact of serial spawning on reproductive allocation. This study investigated whether resource allocation was influenced by parental characteristics and prior spawning history in a serial spawning tropical damselfish (Pomacentrus amboinensis). The offspring attributes of isolated parents of known characteristics were monitored over a 6-week breeding period in the field. Smaller females produced larvae of longer length and larger energy reserves at hatching. This finding is contrary to several other studies that found larger females produce offspring of greater quality. We found that resource allocation in the form of reproductive output was not influenced by the number of spawning events within the breeding season, with larger females producing the greatest number of offspring. Larval characteristics changed as spawning progressed. There was a general decline in length of larvae produced, with an increase in the size of the larval yolk-sac, for all females regardless of size as spawning progressed. This trend was accentuated by the smallest females. This change in larval characteristics may reflect a parental ability to forecast unfavourable conditions as the season progresses or a mechanism to ensure that some will survive no matter what conditions they encounter. This study highlights the importance of accounting for temporal changes in reproductive allocation in studies of reproductive trade-offs and investigations into the importance of parental effects.     

Insights into spawning behavior and development of the European amphioxus (Branchiostoma lanceolatum)

The cephalochordate amphioxus (Branchiostoma sp.) is an important animal model for studying the evolution of chordate developmental mechanisms. Obtaining amphioxus embryos is a key step for these studies. It has been shown that an increase of 3-4 degrees C in water temperature triggers spawning of the European amphioxus (Branchiostoma lanceolatum) in captivity, however, very little is known about the natural spawning behavior of this species in the field. In this work, we have followed the spawning behavior of the European amphioxus during two spawning seasons (2004 and 2005), both in the field and in captivity. We show that animals in the field spawn approximately from mid-May through early July, but depending on the year, they show different patterns of spawning. Thus, even if temperature has a critical role in the induction of the spawning in captivity, it is not the major factor in the field. Moreover, we report some improvements on the methodology for inducing spawning in captivity (e.g. in maintenance, light cycle control and induction of spawning in a laboratory without running sea water system). These studies have important implications for amphioxus animal husbandry and for improving laboratory techniques to develop amphioxus as an experimental animal model.     

Paternal defence,cannibalism and polygamy: factors influencing the reproductive success of painted greenling (Pisces,Hexagrammidae)

The relative costs and benefits of paternal egg-guarding behaviour were identified for painted greenling, Oxylebius pictus, a small fish inhabiting temperate rock reefs. Because males must guard embryos against conspecifics of both sexes and against other fishes, their movements were curtailed, and an energy cost resulted. The males' somatic condition declined in proportion to the time spent guarding. Paternal males partially compensated by cannibalizing a small fraction of their progeny. Male O. pictus thus parasitize females by consuming their spawn. Polygamous matings were nonetheless advantageous for individual females as well as for males. The reproductive success of males and females increased for male broods with greater numbers of clutches, because there was a general dilution of predation loss in larger broods. The success of females, moreover, varied depending on the rank (chronological) order of spawning; success was greater for those that contributed earliest to the male's brood. The results support the hypothesis that the breeding cycles typical of paternal teleosts have evolved partly because of energy costs imposed by site-dependent care. For multiple-clutch species with protracted breeding seasons like O. pictus, the long-term benefits of polygamous spawnings perhaps outweight short-term losses sufficiently for filial cannibalism to evolve and be maintained. Alternatively (or in addition), the foraging milieu may exert a stronger average environmental influence than kin selection does.     

Genetic, spatial, and temporal components of precise spawning synchrony in reef building corals of the Montastraea annularis species complex

When organisms release gametes into the sea, synchrony must be precise to increase fertilization and decrease hybridization. We tagged and genotyped over 400 spawning corals from the three species in the Montastraea annularis species complex. We report on the influence of species, individuals, and genotypes on timing of spawning from 2002 through 2009. During their annual spawning event M. franksi spawns on average 2 h after sunset, whereas M. annularis and M. faveolata spawn 3.5 h after sunset. Only M. franksi and M. annularis have compatible gametes. Individual colonies of the same genotype spawn at approximately the same time after sunset within and across years (within minutes), but different genotypes have significantly different spawning times. Neighboring colonies, regardless of genotype, spawn more synchronously than individuals spaced further apart. At a given distance, clone-mates spawn more synchronously than nonclone-mates. A transplant experiment indicates a genetic and environmental influence on spawn time. There is strong, but not absolute, concordance between spawn time, morphology, and genetics. Tight precision in spawning is achieved via a combination of external cues, genetic precision, and perhaps conspecific signaling. These mechanisms are likely to influence reproductive success and reproductive isolation in a density-dependent manner.     

Coral spawn timing is a direct response to solar light cycles and is not an entrained circadian response

Broadcast spawning corals release gametes into the oceans with extraordinarily accurate timing. While the date of spawning is set by the lunar cycle, the hour/minute of spawning is set by the solar cycle. In this report, we describe experiments that test whether the time of spawning is regulated by an entrained biological clock or whether it is directly controlled by the solar cycle. Montastraea franksi samples were collected on the morning of the predicted spawning. Fragments from colonies were kept under three different lighting conditions and spawning monitored. The three conditions were sunset times of 0, 1 or 2 h earlier than normal. Fragments from the same colony spawned differently under these three conditions, with an early sunset causing a corresponding early shift in spawning. These results indicate that spawn timing is not controlled by a circadian rhythm and that it is directly controlled by local solar light cycle.     

Assessment of field fertilization success in the infaunal polychaete Arenicola marina (L.)

Summary

Previous studies on the fertilization rates in externally fertilizing marine invertebrates have been concerned principally with free spawning epifaunal organisms (e.g., Babcock et al., 1992). A technique has been developed to investigate fertilization success in the intertidal infaunal polychaete Arenicola marina. The spawning period at the East Sands, St Andrews, is predictable from previous data, and this facilitates the study of spawning behaviour. Female worms spawn within their burrows, where the eggs may remain for several days. Male worms release sperm puddles on to the surface of the sediment at low water. The sperm is carried on the incoming tide over the surface of the sediment where it is actively drawn into female burrows by pumping. The decrease in sperm concentration as the tide comes in has been measured, and it is estimated that there are typically about 10⁶ sperm.ml^?1 in the water column and available to the female at 1 min after covering by the incoming tide. By placing the worms into artificial plastic burrows with removable stoppers, they can be transplanted into the field prior to spawning, and removed afterwards whilst retaining all the contents of the tube (worms, gametes, sediment and water).Preliminary data on the fertilization success suggests that it is highly variable (between 0% and 90%), with typical values of 40–60% for Arenicola marina. Laboratory assays investigating fertilization rate with varying sperm concentration have shown that there is a marked increase in success at 10⁴ and 10⁵ sperm per ml, with values of 30–70%. The field data therefore appear to be consistent with laboratory studies.