Environmental and Endocrine Influences on Reproduction of Fundulus heteroclitus

Reproduction in Fundulus heteroclitus is adapted to the temperateclimate and the tidal environment of the coastal marshes whichare its primary habitat. Egg deposition and spawning are closelylinked to the tidal cycle. Concentration of spawning on springtides gives the fish access to sites in the high intertidalzone. Eggs are deposited near the high water mark, usually insand in New England populations and in Spartina alternifloraor empty Geukensia demissa shells in Middle Atlantic and southernpopulations. Aerial incubation of eggs appears to be essentialfor their survival in the salt marsh. Loss of eggs due to lackof fertilization, mortality during development and predationwas always less than 30 percent in our observations. Hatchingoccurs only when the eggs are inundated, usually on spring tides.An endogenous semilunar gonadal cycle appears to be involvedin control of spawning, but an effective environmental synchronizerhas not been found. Cyclic changes in estradiol-17ßaccompany the gonadal cycle, while spawning behavior is stimulatedby neurohypophyseal hormones. Both temperature and photoperiodhave been implicated in the control of seasonal reproductionin F. heteroclitus. Early workers focused on males and concludedthat increasing temperature was the stimulus that initiatedgonadal recrudescence in spring. These experiments were notrigorous tests of the effects of photoperiod. We have shownthat in females both warm temperatures and long photoperiodsare involved. In female F. heteroclitus, as in most mammalsand birds, it is the timing rather than the duration of thelight exposure which determines its effectiveness in stimulatinggonadal maturation. Maintenance of gonadal maturity in femaleF. heteroclitus depends on long photoperiods, but the ovarymay become refractory and regress even when stimulatory photoperiodsare sustained in the laboratory. Ovarian maturity can be stimulatedwith gonadotropin injection in refractory animals, indicatingthat the brain-pituitary axis is the site of refractoriness.The pathway by which light influences ovarian maturity may involvean encephalic photoreceptor, since neither the eyes nor thepineal gland is necessary.     

Waiting for a sign: extended incubation postpones larval stage in the beach spawning California Grunion <Emphasis Type="Italic">Leuresthes tenuis</Emphasis> (Ayres)

Unlike most embryos that hatch on a predetermined timetable, California Grunion Leuresthes tenuis can prolong the embryonic period up to three times longer than the time required for hatching readiness. L. tenuis are teleosts that spawn tidally around the highest spring tides of spring and summer, incubating eggs above the water line. Embryos are competent to hatch in 10 days, however they do not hatch until triggered by an environmental cue, agitation in seawater, as the next spring tides rise. This study examined the growth and survival of L. tenuis embryos and larvae that were all fertilized on the same day, then triggered to hatch after different durations of incubation, up to 35 days post fertilization. L. tenuis embryos that survive extended incubation had decreased yolk reserves and did not advance appreciably in morphological development, even when incubation time was extended to its upper limit. After extended incubation, length of hatchlings was significantly longer than hatchlings from the primary incubation time. Regardless of the duration of incubation, larvae provided food ad libitum grew rapidly and were not significantly different in length at three weeks post hatch. Dry mass increased over time and was not significantly different between larval groups within any post-hatch age. Larval growth and survival after one additional tidal cycle of incubation are not adversely affected, but longer incubation significantly decreases embryonic and larval survival. Large reproductive output, environmentally cued hatching, and plasticity in incubation duration enable L. tenuis to reproduce successfully in the unpredictable sandy intertidal ecosystem.     

SEMI-LUNAR AND LUNAR SPAWNING PERIODICITY IN SOME TROPICAL LITTORINID GASTROPODS

Littoraria strigata, Nodilittorina millegrana and N. pyramidalisall spawned regularly in a rock-flanked Malaysian bay in March-June1985, whereas none had spawned there in July-October 1981. N.millegrana showed the least precise timing with some irregularpreponderance during the days between 2d before and 7d afternew or full moon and their associated spring tides. Spawningof N. pyramidalis was more strictly confined to spring tideswith 90% of eggs appearing between 2d before and 5d after newor full moon. There was no apparent difference between alternatespawnings on new and on full moons. L. strigata also spawnedregularly on the spring tides with 89% of eggs appearing between1 and 5 days after new and full moon. In this case, however,full moon spawnings (when spring tides rose highest) yieldedsome 7 times more eggs on average than the weaker spring tidesat new moons. In laboratory aquaria, the three species all released eggs duringboth night and daytime and through periods of both spring andneap tides with no evidence that periodicity persisted in constantconditions. The different degrees of 2-weekly spawning periodicity, togetherwith littorinids which spawn only monthly, may represent a rangeof adaptation to life at increasing shore heights. (Received 26 November 1985;     

Starvation tolerance of rotifers produced from parthenogenetic eggs and from diapausing eggs: a life table approach

Monogonont rotifers have to cope with environmental variationby producing diapausing stages. Cyclically, parthenogeneticrotifers produce females by (i) parthenogenetic eggs and (ii)sexually produced eggs. Parthenogenetic eggs hatch shortly afterovoposition, frequently while they are carried by their mothers.Sexual eggs go into dormancy and hatch in the sediment, in responseto specific environmental cues. Therefore, it is expected thatpost-diapause and parthenogenetic offspring would face differentenvironments. Moreover, resource allocation is higher in diapausingeggs than in parthenogenetic eggs. In this study, the responseto starvation of females obtained from parthenogenetic eggsand from diapausing eggs of the rotifer Brachionus plicatiliswas compared. Starved individuals showed unexpectedly long maximumlongevity (12 days). The average lifespan and the lifespan varianceof individuals hatched from parthenogenetic eggs were higherthan those of individuals hatched from diapausing eggs. Contrastingwith a previous experimental finding, our results do not supportthe hypothesis that high resource allocation in diapausing eggscauses high starvation tolerance. Lower survival of individualshatched from diapausing eggs could be caused by the diversionof energy to other functions (i.e. maintenance during diapauseor the hatching event), or the result of the variance in theallocation of resources in parthenogenetic eggs. We hypothesizethat resource allocation in diapausing eggs follows a conservativepattern, while it is more opportunistic in parthenogenetic eggs. This paper was presented at Plankton Symposium III, held atFiguera da Foz, Portugal between 17 and 20 March 2005, underthe auspices of the University of Coimbra and the Universityof Aveiro, and coordinated by Mário Jorge Pereira andUlisses M. Azeiteiro.     

THE LIFE-CYCLE AND REPRODUCTIVE ACTIVITY OF ACHATINA FULICA (BOWDICH) IN LABORATORY CULTURE

The life-cycle, growth characteristics and reproductive activityof Achatina fulica were studied in laboratory culture. Six generationshave been completed. Sexual maturity is reached as early as5 months of age, with a peak in egg production between 210–270days. Egg clutches contain 100–200 eggs which hatch synchronouslywithin a 24-hour period. The incubation period (days betweenoviposition and hatching) was determined exactly. The rangeof incubation times was 1–25 days, due to a variable degreeof ovoviviparity exhibited by the snail. The success of laboratoryculture provides a basis for experimental studies in developmentalneurobiology utilizing Achatina fulica. ^*Present address and to whom correspondence should be addressed:Jerry Lewis Neuromuscular Research Center, UCLA School of Medicine,Los Angeles, California 90025 U.S.A. (Received 14 February 1983;     

Influence of Inoculum Density,Host, and Low-temperature Period on Delayed Hatch of Meloidogyne javanica Eggs

Most eggs of M. javanica hatch within several days when incubated in water. Those that do not are said to show delayed hatching. Several experiments were conducted to determine the effect of specific conditions on the percentage of eggs with delayed hatch. Six initial inoculum densities ranging from 100 to 20,000 eggs per pot did not influence egg hatch within a 45-day incubation period. In a 60-day test, the percentage of eggs hatching after more than 20 days was low for egg masses removed from carrot and okra and high for those from pepper and bean. Increasing exposure to cold temperature (8 C) from 7 to 30 days tended to delay hatch.     

Thermohaline tolerance and embryonic development in capelin eggs (Mallotus villosus) from the Northeast Atlantic Ocean

Capelin (Mallotus villosus) displays alternative reproductive modes throughout its circumpolar distribution. This predicts divergent thermohaline tolerance of eggs because they are incubated in either a steady offshore or variable intertidal environment. We investigate herein thermohaline tolerance of eggs from the offshore spawning Barents Sea capelin. Subsequently, we compare our data with those previously published on other offshore and intertidal spawning capelin populations across the Northeast Atlantic Ocean, with the aim of determining possible patterns in the thermohaline tolerance of eggs from the alternative reproductive modes. In a 2?×?4 factorial design various combinations of salinities and temperatures had only negligible effect on the survival of eggs until first hatch. The embryonic development rate from fertilisation until first hatch across populations and between the two reproductive modes suggested non-local thermohaline tolerance towards the physical factors during development. Finally, no differences were observed in salinity tolerance from fertilisation to first hatch between populations representing different reproductive modes. The present findings demonstrate wide thermohaline tolerance of capelin eggs regardless of population origin and reproductive mode.     

Semi-monthly reproductive cycles in male and female fiddler crabs, Uca pugnax

In the fiddler crab, Uca pugnax, numbers of displaying males, numbers of burrows with half-dome superstructures (hoods), numbers of females with decalcified vulvar opercula, and numbers of females hatching their eggs, all follow a semi-monthly cycle with peaks near the time of the spring tides. These are all aspects of reproduction in Uca. Decalcification of female opercula, although necessary for mating and egg deposition, is neither necessary nor sufficient for behavioural response to courting males. In fact, behavioural responsiveness to males probably precedes decalcification in most cases. The different aspects of the reproductive cycle are coordinated so that females and males are ready to mate at about the same time near one spring tide, and the resulting eggs will be ready to hatch on the next spring tide. Two decalcified females which were kept isolated from males produced fertile clutches, suggesting that stored sperm from a previous mating may fertilize eggs in a later cycle. This may be one reason for the relative rarity of observed matings or precopulatory behaviour in Uca.     

The Role of Fundulus heteroclitus in Salt Marsh Trophic Dynamics

The common mummichog, Fundulus heteroclitus, functions as bothpredator and prey in the trophic structure of east coast tidalmarshes. Although mummichogs are generally considered importantto energy transformations within marshes, few studies have convincinglydemonstrated that predation by F. heteroclitus affects the abundanceof salt marsh benthic invertebrates. Thus far investigationsof this type have dealt only with the direct effects of adultmummichogs. The results of recent experiments have suggestedthat by controlling smaller predators, mummichogs may indirectlyhave a positive effect on the densities of some infaunal marshinvertebrates. Our current knowledge of larval and juvenilemummichogs in their natural habitat is minimal. Unlike the adults,which can utilize the intertidal zone only when it is flooded,the young remain on the marsh even at low tide, inhabiting shallowpuddles of residual tidal water that form between clumps ofvegetation and around fiddler crab (Uca sp.) burrows. The importanceof F. heteroclitus in salt marsh communities will remain incompletelyunderstood unless future studies consider the role of larvaland juvenile mummichogs. Although many species of fishes andwading birds feed on mummichogs, the blue crab (Callinectessapidus) is probably the major predator of adult F. heteroclitusin the intertidal salt marsh. Predation by adult mummichogsand xanthid crabs (e.g., Eurytium limosum) may contribute tothe high mortality of larval and juvenile Fundulus.     

The effect of reproductive condition on egg production rates in the planktonic copepod Centropages typicus

Centropages typicus females oscillated between a dark, ripestage of oogenesis and a light, unripe stage of primary oocytes.The number of eggs released after 24 h under different experimentalconditions was strongly dependent on the reproductive statusof females at the time of incubation.     

Behavioral and Morphological Adaptations by Galapagos Land Iguanas (Conolophus subcristatus) to Water and Energy Requirements of Eggs and Neonates

We examine the importance of both the hydric environment ofnaturally incubating reptilian eggs and the energetic needsof hatchlings via an investigation of reproduction in Galapagosland iguanas (Conolophus subcristatus). Hatching success ofegg clutches and the size of subsequent hatchlings are bothpositively correlated with the water potential of natural nests,as predicted from previous laboratory experiments. Water potentialsrepresenting optimal incubation environments are available foronly a brief period in nature, and depend upon the same seasonalrainfall as does food abundance for the emerging hatchlings.The temporal placement of the reproductive season of Conolophussubcristatus balances these conflicting needs for water by eggsand energy by hatchlings. Oviposition occurs slightly beforesuitable water potentials are reached and hatchling emergenceoccurs after the peak in food abundance. Morphological adaptations by Conolophus subcristatus to theirprecarious reproductive phenology include greater amounts ofalbumen in their eggs, and greater energy reserves in emergenthatchlings than most other lizard species. These adaptationslessen the severity of an arid environment where water becomesavailable for periods too short to allow both oviposition andhatching to be temporally placed in an optimal manner.     

Reproductive performance of the desert pupfish (Cyprinodon n. nevadensis) in relation to salinity

Synopsis The reproductive performance of the desert pupfish, Cyprinodon n. nevadensis, was tested by exposing breeding pairs to a range of salinities from 0.1 to over 40 In terms of eggs g^–1 body weight day^–1 , eggs per spawning and embryo viability. reproductive performance was optimal at 10 Performance decreased above and below 11 producing a plateau of statistically equivalent reproductive performance from 0.2 to 20 The upper salinity limit placed on reproduction is the tolerance of the eggs, since a few eggs are laid above 20but they do not hatch. These small fish probably never experience the upper salinity limit for reproduction in their native habitat. The tolerance limits for reproduction are compared with a lethal range from less than 0.1 to 53in a 96 h LD-50 test.     

REPRODUCTIVE CYCLE AND FECUNDITY OF THE DATE MUSSEL LITHOPHAGA LITHOPHAGA (BIVALVIA: MYTILIDAE)

The reproductive cycle and fecundity of the date mussel Lithophagalithophaga, L. a well-known, ediblespecies has been examined.Sexes are separate and the mean number of eggs per each gonadis 1.894 x 10⁶ ± 1.044 x 10⁶ S.D. Reproduction firstoccurs at an age of 2⁺years and at a shell length greater than0.9 cm. Gonad activity is annual and is observed at all ages.The sex ratio for individuals up to 7 cm is 3:1 in favour ofthe males, whereas it becomes 1:1 for individuals greater than7 cm. The release of gametes by males and females occurs almostsimultaneously and begins immediately after a decline in thehighest water temperature ( 27°C), an increase in salinity(>31) and a decrease in the dissolved oxygen (6.5 ppm). Smallpercentages of mature individuals appear during the first wintermonths thus lengthening the reproductive period. This phenomenonis attributed to the temperature difference in deeper waters,the delay in gamete release by young individuals, tide, waveaction and changes in salinity. The fecundity of Lithophagalithophaga is high because its life-cycle is adversely affectedby environmental factors such as waves and tides. Fecundityincreases with shell length, more so with total wet weight andmainly with the age of the animals. Summer seems to be the suitableseason to exploit the date mussel of shell lengths > 5 cm. (Received 6 September 1993; accepted 10 March 1994)     

Quantification of Reproductive Isolating Barriers Between Two Naturally Hybridizing Killifish Species

Understanding the relative importance of various reproductive barriers to the early stages of speciation is an essential question in evolutionary biology. The closely related killifishes Fundulus heteroclitus and F. grandis occasionally hybridize in a small region in coastal Northeastern Florida showing that while barriers to reproduction exist, they are incomplete. The objective of this study was to elucidate barriers to reproduction between F. heteroclitus and F. grandis in the lab, as well as to quantify their strengths and relative contributions to reproductive isolation. Pre-zygotic (mating and fertilization) and post-zygotic (hatching) barriers were investigated by performing a variety of choice and no-choice laboratory mating experiments. Under no-choice conditions, barriers to mating had the greatest influence on hybrid production in F. grandis, whereas hatching barriers contributed to the majority of reproductive isolation in F. heteroclitus. Under choice conditions, however, pre-zygotic barriers had the greatest influence on hybrid production in both species. The total reproductive isolation that was observed in females of each species was stronger in F. heteroclitus than in F. grandis, and was nearly complete in F. heteroclitus females under choice conditions and was of moderate strength in F. grandis females. These results reveal an asymmetry in the potential gene flow between these two species, with F. grandis being more likely to hybridize than F. heteroclitus in the absence of environmental influences. No-choice backcrosses were also conducted and showed that at least some F1 hybrids are fertile. The observation that pre-zygotic barriers tend to be stronger than post-zygotic barriers in the early stages of speciation is consistent with similar studies in other organisms.     

Zooplankton and ichthyoplankton communities in a temperate estuary: spatial and temporal patterns

Zooplankton and ichthyoplankton assemblages were studied fromJanuary 2003 to June 2004 in a temperate shallow estuary (Mondegoestuary, Portugal). Monthly sampling was performed at five stationsat high and low tides, with subsurface tows with 335 and 500µm mesh Bongo nets. Analysis of variance (ANOVA) showeda significant effect (P < 0.05) of the mesh size of the neton the abundance of main zooplankton groups. On average, theabundance of the 500 µm taxocenosis was 67 and 102 timeslower than the 335 µm taxocenosis at high and low tidesrespectively, especially in the south arm. More than 80 specieswere identified in the zooplankton samples. The upper reachesof northern arm are dominated by freshwater crustacean mesozooplanktonlike Acanthocyclops robustus and Diaptomus spp. and the cladoceransDaphnia, Ceriodaphnia and Bosmina, often being codominant. Inthe southern arm, the resident estuarine copepod Acartia tonsawas dominant, eventually being the most abundant taxon. Marinereaches of estuary are usually dominated by the marine calanoidcopepods Acartia clausi and Temora longicornis and the siphonophoresMuggiaea atlantica. Concerning the ichthyoplankton, this wasdominated by the larvae of estuarine resident species, mainlyPomatoschistus sp., and eggs of Engraulis encrasicolus. Theabundance of Pomatoschistus sp. larvae was positively correlatedwith water temperature. Statistical analysis (canonical correspondenceanalysis) used to determine the spatiotemporal structure ofthe zooplankton assemblages and its correlation with environmentalvariables showed that salinity and temperature were the mainfactors influencing the distribution of zooplankton. The resultsobtained also showed that abundance was strongly influencedby the hydrological circulation pattern and direct or indirecthuman impacts that occur in each arm of the estuary. This article was presented at Plankton Symposium III, held atFiguera da Foz, portugal, between 17 and 20 March 2005, underthe auspices of the University of Coimbra and the Universityof Aveiro, and coordinated by Mário Jorge Pereira andUlisses M. Azeiteiro.     

Clutch detachment in a copepod after capture by a predator

When the egg-carrying copepod, Eudiaptomus gracilis, is capturedby larvae of predatory Chaoborus species (C.obscuripes or C.flavicans)in laboratory experiments, the external egg clutch detachesin 16-29% of caw, thereby allowing the eggs to escape from ingestion.Clutch detachment may be due to active removal by the femalebefore she is eaten or the result of accidental loss by thepredator during handling. Detached clutches were significantlylarger than thase that were ingested along with the female.Clutch detachment is advantageous to the female since ingestedeggs are lost, whereas detached eggs hatch normally and maypotentially propagate the genotype. Chaoborus predation maytherefore influence the evolution of clutch size and the degreeof iteroparity in Eudiaptomus.     

Influence of temperature on hatching of winter eggs of fruit-tree red spider mite, Panonychus ulmi (Koch)

The effects of the duration and degree of chilling, and the temperature of incubation, on hatching of winter eggs of Panonychus ulmi (Koch) were investigated. For chilling, 0°C and 5°C were more effective than — 5° and 9°, and the limits for the reaction were close to — 10° and 15°. As the chilling period was increased from 60 to 200 days, the percentage hatch on incubation at 21° increased, and the mean incubation time and its variance decreased. Before the maximum effect of chilling was achieved, percentage hatch on incubation at 9° and 15° was higher than at 21°; 27° was lethal to most winter eggs though not to summer eggs. After chilling, the later stages of diapause development could occur at temperatures from 0° to 21°₎ i.e. above and below the threshold temperature for morphogenesis, 6–7° in both winter and summer eggs. Diapause development cannot, therefore, be a unitary process. The significance of the results is discussed in relation to forecasting the time of hatch in the field, and to the phenological aspects of hatching in the spring.     

Climate and geographic trends in hatch delay of the treehole mosquito,Aedes triseriatus Say (Diptera: Culicidae)

Eggs of Aedes triseriatus mosquitoes are stimulated to hatch when inundated with water, but only a small fraction of eggs from the same batch will hatch for any given stimulus. Similar hatching or germination patterns are observed in desert plants, copepods, rotifers, insects, and many other species. Bet hedging theory suggests that parents stagger offspring emergence into vulnerable life history stages in order to avoid catastrophic reproductive failures. For Ae. triseriatus, a treehole breeding mosquito, immediate hatching of an entire clutch leaves all of the parent's progeny vulnerable to extinction in the event of a severe drought. Natural selection has likely favored parents that pursued a bet hedging strategy where the risk of reproductive failure is distributed over time. Considering treehole mosquitoes, bet hedging theory could be used to predict that hatch delay would be positively correlated with the likelihood of drought. To test this prediction, we collected Ae. triseriatus from habitats that varied widely in mean annual precipitation and exposed them to several hatch stimuli in the laboratory. Here we report that, as predicted, Ae. triseriatus eggs from high precipitation regions showed less hatch delay than areas of low precipitation. This strategy probably allows Ae. triseriatus to cope with the wide variety of climatic conditions that it faces in its extensive geographical range.     

Artificial incubation of trumpeter swan eggs: Selected factors affecting hatchability

Eggs collected from captive trumpeter swans (Cygnus buccinator) in 1993 (n = 33) and 1994 (n = 42) were artificially incubated with careful monitoring to identify factors contributing to the low hatch success reported by the Ontario Trumpeter Swan Restoration Program. Fertility was > 80% in both years, whereas hatch success of fertile eggs was 14.3% (n = 4) of 28 eggs in 1993 and 37.1% (n = 13) of 35 eggs in 1994. Necropsy of non‐viable eggs indicated a high incidence of embryonic mortality during early and late incubation. Early embryonic mortality was associated with egg storage times exceeding 7 days (P < 0.05) and bacterial contamination of eggs (P < 0.01). Late mortality was associated with (P < 0.001) increased weight loss during incubation period and may have resulted from incubator temperature and humidity fluctuations. We established patterns of weight loss for eggs and determined that hatched eggs lost 11–15% of initial mass and that weight loss >15% resulted in embryo mortality. Results from this study indicate that collection and handling of eggs before incubation and precise control of the incubator environment are critical to hatchability of eggs. Zoo Biol 18:403–414, 1999. © 1999 Wiley‐Liss, Inc.     

Embryonic Development and Rates of Metabolic Activity in Early and Late Hatching Eggs of the Major Malaria Vector Anopheles gambiae

Anopheles gambiae eggs generally hatch at the completion of embryo development; two-three days post oviposition. However, staggered or delayed hatching has been observed whereby a single batch of eggs shows marked variation in time-to-hatch, with some eggs hatching 18 days post oviposition or later. The mechanism enabling delayed hatch has not been clearly elucidated but is likely mediated by environmental and genetic factors that either induce diapause or slow embryo development. This study aimed to compare metabolic activity and embryonic development between eggs collected from sub-colonies of the baseline Anopheles gambiae GAH colony previously selected for early or late time-to-hatch. Egg batches from early and late hatch sub-colonies as well as from the baseline colony were monitored for hatching. For both time-to-hatch selected sub-colonies and the baseline colony the majority of eggs hatched on day two post oviposition. Nevertheless, eggs produced by the late hatch sub-colony showed a significantly longer mean time to hatch than those produced by the early hatch sub-colony. The overall proportions that hatched were similar for all egg batches. CO₂ output between eggs from early and late hatch sub-colonies showed significant differences only at 3 and 7 days post oviposition where eggs from the early hatch and the late hatch sub-colony were more metabolically active, respectively. No qualitative differences were observed in embryo development between the sub-colonies. It is concluded that all viable embryos develop to maturity at the same rate and that a small proportion then enter a state of diapause enabling them to hatch later. As it has previously been shown that it is possible to at least partially select for late hatch, this characteristic is likely to involve genetic as well as environmental factors. Delayed hatching in An. gambiae is likely an adaptation to maximise reproductive output despite the increased risk of desiccation in an unstable aquatic environment.