Effect of Eyestalk-Ablation on Circulating Ecdysteroids in Hemolymph of Snow Crabs, Chionoecetes opilio: Physiological Evidence for a Terminal Molt

Bering Sea snow crabs (Chionoecetes opilio) are a commerciallyimportant crab harvested in the Bering Sea. Optimal managementof this species requires an understanding of the biology ofthis crab that is currently incomplete. Fisheries managers applya continuous growth model in their management of snow crab,which assumes that male crabs increase in size throughout theirlifespan. Male snow crabs undergo a morphometric molt that leadsto a disproportionate increase in chelae size and it is stilldebated whether this molt is associated with a terminal molt.This study was conducted to determine whether adult male C.opilio are anecdysic. Using current knowledge of the hormonalregulation of crustacean growth, snow crab physiology was manipulatedto induce an increase in molting hormones (ecdysteroids). Sincefemale snow crabs are known to undergo a terminal molt afterattaining reproductive maturity, we compared ecdysteroid levelsin eyestalk-ablated terminally molted females, small-clawedmales and large-clawed males. Snow crabs were collected fromthe Bering Sea and maintained in circulating seawater at approximately6°C. Animals were either eyestalk-ablated or left intact.Ecdysteroid levels in hemolymph were quantified using an enzyme-linkedimmunosorbant assay (ELISA). Circulating ecdysteroids were significantlyhigher in small-clawed male crabs when compared to large-clawedmales or terminally molted females. Eyestalk-ablation increasedcirculating ecdysteroids in small-clawed males, but had no significanteffect on circulating ecdysteroids in large-clawed males orin terminally molted females.     

A Review of Size at Maturity in Male Tanner (Chionoecetes bairdi) and King (Paralithodes camtschaticus) Crabs and the Methods Used to Determine Maturity

This paper reviews existing studies on the size of sexual maturityfor male Tanner or snow crab (Chionoecetes bairdi), a brachyuran,and the anomuran red king crab (Paralithodes camtschaticus).In this report the term sexual maturity is denned as the abilityto reproduce. A variety of indirect and direct methods thathave been used to determine maturity are reviewed. Examiningthe vas deferens for the presence of spermatophores was usefulin determining the size at which males first become mature.Breeding experiments in the laboratory demonstrated that mostmales, from both species, that produced spermatophores couldbreed with soft-shelled mates. Males of both species can breedat smaller sizes than do females. Morphometric techniques basedon reproductive tract weights and chela morphometry overestimatedthe sizes at which males mature in both species. Previous experimentsfor Tanner crab, which have internal fertilization, suggestsmall mature males can fertilize two to five females. Breedingexperiments showed recently matured red king crab do not appearto be able to fertilize more than one female per breeding season,while males nearing harvestable size can fertilize more thanone female. Breeding experiments and in situ observations of grasping pairsappear to be the most feasible methods for identifying malesize at maturity for these species. The value of morphometricestimations for determining when males mature is questionable.     

Larval release in a sub-euphotic zone invertebrate triggered by sinking phytoplankton particles

This paper reports field results showing that the larval releaseof the snow crab, Chionoecetes opilio, in a subarctic fjordis phased with the sedimentation peak of phytoplankton particlesoriginating from the surface spring bloom, but with a lag of2–3 weeks. Further laboratory experiments with egg-bearingfemales showed a significant induction of larval release whenthey were exposed to senescent cultures of different phytoplanktonspecies, as opposed to controls and females exposed to freshcultures. These results are interpreted and discussed in thecontext of possible strategies evolved by the snow crab to enhanceits reproductive success. The field and laboratory evidencepresented here represents the first instance indicating thatphyto-detritus can act as a chemical cue for initiating andsynchronizing reproductive activities in sub-euphotic invertebrates.Selection for control of larval release by downward flux ofalgal particles may be especially important for invertebratepopulations in high-latitude areas, where the development ofplankton communities shows a high level of seasonality.     

The influence of insect juvenile hormone agonists on metamorphosis and reproduction in estuarine crustaceans

Comparative developmental and reproductive studies were performedon several species of estuarine crustaceans in response to threejuvenile hormone agonists (pyriproxyfen, methoprene and fenoxycarb).Larval development of the grass shrimp, Palaemonetes pugio,was greater than two orders of magnitude more sensitive to disruptionby methoprene and fenoxycarb than was embryonic development.Developing larvae of the mud crab, Rhithropanopeus harrisii,exhibited reduced metamorphic success at lower concentrationsof methoprene and pyriproxyfen than grass shrimp larvae. Theseresponses suggest that the more rigidly controlled metamorphicprocess in crabs is more sensitive to compounds acting as endocrinedisruptors than is the more flexible metamorphic pattern inshrimp. The final crab larval stage, the megalopa, was moresensitive to methoprene and fenoxycarb exposure than earlierzoeal stages. Mud crab larvae exposed to fenoxycarb had reducedbiomass and lipid content, particularly triglycerides and sterols.Concentrations of fenoxycarb which reduced the reproductivecapacity in single life-cycle exposures of the estuarine mysid,Americamysis bahia, were similar to those concentrations whichinhibited metamorphosis in grass shrimp. Juvenile mysids releasedby exposed adults and reared through maturation without furtherexposure produced fewer young and had altered sex ratios (lowerpercentages of males) at lower parental-exposure concentrationsthan directly affected parental reproduction. These transgenerationalresponses may well be a product of irreversible effects duringdevelopmental exposures which become apparent following maturationand initiation of reproduction. These findings support usinga functional approach as an appropriate screening procedureto evaluate potential environmental endocrine-disrupting chemicalsin aquatic environments.     

Reproductive parasitism of broodcare helpers in a cooperatively breeding fish

Large male helpers in the cooperatively breeding cichlid Neolamprologus pulchergain reproductive success by parasitizing the reproductive effort ofmale territory owners. Under controlled, experimental conditionswe examined the genetic relatedness between the members of broodpairs (n = 14), their male helpers (n = 8), and offspring (n= 292) in seven families. We used multilocus DNA fingerprintingto check for potential reproductive parasitism by male helpersand to assess their fertilization success. Of offspring producedin these families, 10.3% were sired by helpers. In parasitizedbroods, helper fertilization success varied between 12.5% and35.8%. Male helpers parasitized parental reproduction when theirbody size exceeded 4.5 cm standard length (SL), even thoughsexual maturity may be reached much earlier (3.5 cm SL). Twoof three parasitic helpers were punished by severe aggressiveattacks when parasitizing the reproduction of breeders, whichled to their expulsion from the territory. This study demonstratesa potential fitness benefit to broodcare helpers that is oftenneglected. It also points to the delicate balance that may exist betweencooperative and competitive behavior in cooperatively breeding species.     

Male-caused failure of female reproduction and its adaptive value in alpine marmots (Marmota marmota)

We studied reproductive performance of free-living alpine marmots (Marmotamarmota) for 14 years in the National Park of Berchtesgaden, Germany.Female reproduction was influenced by body condition and social factors.Reproduction depleted fat reserves, and only females emergingfrom hibernation with sufficient body mass were able to reproducesuccessfully. Marmots lived in social groups in territoriesdefended by a dominant male and female. Subordinate femalesnever reproduced, regardless of body mass. Territory takeoversby males impaired reproduction of dominant females, but onlyif the takeover occurred after the mating period. Reproductivefailures occurred despite clear signs of pregnancy such enlargednipples or late molt. Decreasing progesterone levels after themating period and the lack of evidence for direct infanticideby new territorial males suggest a block of pregnancy as a likelyexplanation for reproductive failures in groups with male takeoversduring gestation. Rendering female reproduction impossible increasedfuture reproductive success of new territory owners. Nonparous femalessaved the energetic cost of maternal investment and thus emergedwith higher body mass in the following spring. In line withthis, females failing to wean young had higher reproductivesuccess in the subsequent year.     

First observations of temperature tolerances of adult male snow crab (Chionoecetes opilio) from the Barents Sea population and the effects on the fisheries strategy

This study investigates the effects of temperature on the survival, food intake, oxygen consumption and growth during long-term live holding of captive male snow crab (Chionoecetes opilio) (average?=?0.7?kg). The crabs were held at three different temperatures, 3, 6 and 9°C. The trials were done using groups of snow crabs held in nine land-based holding tanks (three replicates per temperature treatment). The results showed that temperature had a significant effect on survival. The survival rate at 3°C (61%) was significantly higher than at 6°C (33%) and at 9°C (28%). Specific oxygen consumption rates of unfed crab at 6°C were significantly higher than at 9°C and 3°C. In summary, the current study shows that the Barents Sea snow crab have a narrow temperature range in which they thrive compared with the Barents Sea red king crab (Paralithodes camtschaticus). Barents Sea snow crab has similar metabolic and physiological attributes to other major snow crab populations. In conditions when ambient temperatures are approximately 6°C, it may prove beneficial for animal welfare and also be financially advantageous to reduce ambient water temperatures in live snow crab holding facilities on boat or on land.     

Effects of social structure on reproductive activity in male fathead minnows (Pimephales promelas)

The selection of alternative reproductive phenotypes is oftenthought to be the result of physiological state, with smallindividuals forced energetically to postpone the allocationof resources to reproduction. However, for male fathead minnows(Pimephales promelas), we show that seasonal reproductive activityis modulated by social status. In enclosure and pond experiments,small males advanced their reproductive condition, held nesting territories, and spawned earlier in the reproductive seasononly when large males were absent or removed from the population.Since differences in the timing of reproduction among smallmales were not size- or condition-dependent, the common explanationfor the selection of alternative reproductive phenotypes, basedon state-dependence, is insufficient. In the absence of large,socially dominant individuals, small males produced comparablenumbers of offspring as the treatment with large males, although the offspring of these uninhibited small males were smallerat the end of the growing season than the young of large males.Thus, interactions among conspecifics may account for muchof the phenotypic diversity observed within and among naturalfathead minnow populations, through their direct and indirecteffects on growth, recruitment and survival.     

Assessing Costs of Reproduction in Mountain Birch: The Importance of Considering the Modular Level

We studied the effects of production of male catkins on growthand the subsequent year's male catkin production in mountainbirch, Betula pubescens subsp. czerepanovii at both the shootand branch level. (Shoot is defined here as the product of asingle growing season while branch refers to a structure consistingof several shoots.) Like heterophyllous trees in general, mountainbirch canopy expansion takes place via the production of longshoots. We found phenotypic trade-offs between long shoot growthand male reproduction at the shoot level in the year of catkinproduction. Generative parental long shoots (long shoots withmale catkins) were significantly shorter than the vegetativeones (long shoots without a male catkin). In contrast, we foundno effects of male reproduction on the subsequent year's malecatkin production at the shoot level. Although the mean lengthof secondary long shoots (long shoots growing from the lateralbuds of parental long shoots) did not differ between vegetativeand generative parental long shoots, there was considerablebetween-individual variation in the response of individual trees.In addition, production of male catkins diminishes canopy expansionin mountain birch because the number of secondary long shootsproduced by generative parental long shoots was smaller thanthat of vegetative parental long shoots. At the branch level,the association between total long shoot growth and male catkinproduction was positive, i.e. no trade-off was found. This maybe because the strong sink strength (the ability of a branchto import assimilates from elsewhere in the tree) of branchesbearing reproductive long shoots masks possible trade-offs.We emphasize the importance of considering several levels ofthe modular hierarchy when analysing costs of reproduction inmodular organisms. Copyright 2000 Annals of Botany Company Hierarchical structure, modularity, mountain birch, short and long shoots, trade-off, Betula pubescens subsp.czerepanovii (Orlova) Hämet-Ahti     

Reproductive Inhibition of Female Peromyscus leucopus: Female Competition and Behavioral Regulation

Maturational delay of young female mice as the result of exposureto grouped female odors and reproductive inhibition as the resultof exposure to isolated adult females have both been observed(Drickamer, 1974; Skryja, 1978). Each has the potential to reducethe growth rate of populations. Reductions in a female's reproductionfacilitated by social stimulation from other females, whileeffective in reducing population growth, may in the case, ofmaturational delay and reproductive inhibition be an epiphenomenonor exadaptation of selection for improved relative reproductivesuccess in the females possessing these abilities. The ultimateoutcome of these selective processes may be the buffering ofpopulation numbers, but the selective forces may operate tomaximize a female's relative reproductive success. A females'relative reproductive success can be maximized by either increasingher own reproduction or decreasing the reproductive output ofother females. A body of evidence exists to suggest that inPeromyscus mamculatus and Peromyscus leucopus, females are physiologicallyconstrained and unable to increase their own reproduction. Ifthe assumption of physiological restraint is correct, then themost effective way for females to maximize their relative reproductivesuccess is to reduce the reproductive output of their competitors.In this paper, maturational delay and reproductive inhibitionas they affect both the adult female and young females are discussed.Examination of these effects reveals that while they can beeffective in population regulation, their main function andthe selective process that produced them is at the level ofindividual reproduction.     

Reduced reproductive effort in male field crickets infested with parasitoid fly larvae

Some populations of the field cricket Teleogryllus oceanicusare parasitized by the phonotactic fly Ormia ochracea. Flieslocate crickets by their song and deposit larvae onto them.The larvae develop inside the cricket for 1 week before killingthe host upon emergence. The reproductive compensation hypothesispredicts that parasitized crickets should increase their reproductiveeffort during the initial stages of infestation to offset theloss of fitness resulting from their shortened life span. An alternative hypothesis predicts that parasitized crickets willdecrease reproduction, either because they are unable to reproduceor because selection acting on the parasitoid favors decreasedhost reproduction. In laboratory experiments, parasitized malecrickets had reduced reproductive effort (spermatophore production,calling, mating activity, and mass allocated to reproductivetissue) compared to unparasitized males. Parasitized males fedad libitum showed no evidence of allocating a greater proportionof their resources to reproduction. Parasitized and healthymales did not differ significantly in resting or maximal metabolicrates, although this may have been due to the substantial contributionof larval respiration to the metabolic rate of the host—parasitoidcomplex. These results are consistent with previous studiesand suggest that T. oceanicus males parasitized by O. ochraceado not increase their reproductive effort. We discuss potentialreasons that crickets do not increase reproductive effort inresponse to fly larvae and address difficulties in demonstratingaltered life-history patterns in response to parasitism.     

Reproductive skew among males in a female-dominated mammalian society

The purpose of this study was to document patterns of reproductiveskew among male spotted hyenas (Crocuta crocuta), a speciesin which many normal mammalian sex roles are reversed. We usedpaternity determined from 12 microsatellite markers togetherwith demographic and behavioral data collected over 10 yearsfrom a free-living population to document relationships among reproductive success (RS), social rank, and dispersal statusof male hyenas. Our data suggest that dispersal status andlength of residence are the strongest determinants of RS. Natalmales comprise over 20% of the adult male population, yet theysire only 3% of cubs, whereas immigrants sire 97%. This reproductiveadvantage to immigrants accrues despite the fact that immigrants are socially subordinate to all adult natal males, and it providesa compelling ultimate explanation for primary dispersal inthis species. High-ranking immigrants do not monopolize reproduction,and tenure accounts for more of the variance in male reproductivesuccess than does social rank. Immigrant male hyenas rarelyfight among themselves, so combat between rivals may be a relativelyineffectual mode of sexual selection in this species. Instead,female choice of mates appears to play an important role in determining patterns of paternity in Crocuta. Our data supporta "limited control" model of reproductive skew in this species,in which female choice may play a more important role in limitingcontrol by dominant males than do power struggles among males.     

Metamorphosis season from megalopa to the first crab stage in snow crab Chionoecetes opilio and red snow crab C. japonicus (Crustacea,Decapoda, Majoidea) in the Sea of Japan,estimated from captive culture

ABSTRACT

Metamorphosis season of megalopae to the first crab stage in snow crab Chionoecetes opilio and red snow crab C. japonicus was inferred by culturing wild-born megalopae collected from the Sea of Japan. Metamorphosis occurred from late June to late July (mainly in July) in snow crab, and from early July to early October (mainly from August to September) in red snow crab. The number of days required from the time of collection to metamorphosis was less than the intermoult period previously reported for snow crab megalopae. However, the developmental period of the megalopae was estimated as substantially longer in red snow crab than in snow crab. Previous studies have shown that the hatching season and the period of the zoeal stage in both two species are similar. These results suggest that a different metamorphosis season between the two species would be due to a difference in their megalopal intermoult period.     

Characterization of enzymatic hydrolyzed snow crab (Chionoecetes opilio) by-product fractions: A source of high-valued biomolecules

Snow crab (Chionoecetes opilio) constitutes valuable and nutritional sources of components, such as proteins, lipids and chitin. The present investigation was undertaken to evaluate the feasibility of applying a pilot scale enzymatic hydrolysis process of snow crab by-products, followed by fractionation, in order to recover enriched high-valued compounds. The yield of snow crab by-products recovered after manual processing; on a dry weight was 87.4%. The by-products (raw materials) were mainly moist (approximately 78%), and contained 42.9% proteins, 14.8% lipids, 25.7% minerals, 16.2% chitin, all expressed on a dry weight. The fatty acid profile of snow crab by-products and all fractions obtained following processing showed a higher content in mono-unsaturated fatty acids (MUFAs; approximately 50%), followed by polyunsaturated fatty acids (PUFAs; approximately 20%) and saturated fatty acids (SFAs; approximately 15%). The n − 3/n − 6 ratio was approximately 10 and represents a good index of nutritional value for snow crab oil by-products. Most protein enriched fractions demonstrate a well-balanced amino acid composition, notably the most essential amino acids. These protein fractions are characterized by biomolecules having a relatively low molecular weight (35 kDa and less) range. The enzymatic hydrolysis process developed in this study shows that snow crab by-products should to be viewed as having the potential of being identified as high-valued products. Even though the process could be optimized, it is controllable, and depending on hydrolyses conditions, the products obtained are reproducible and well defined. Results presented in this study indicate that snow crab by-products may serve as excellent nutritional components for future applications in the health and food sectors.     

Moulting and growth in earlier and later moulters of adolescent male snow crabs (Chionoecetes opilio) (Brachyura: Majoidea) under laboratory conditions

Growth and moulting of male adolescent snow crabs were investigated in a laboratory culture experiment. The moulting season of adolescents was separated into two groups of earlier and later, and they were regarded as normal and skip moulters, respectively. The existence of male snow crab skip moulters was first demonstrated in the Sea of Japan. Although the precise moulting season of the earlier moulter could not be revealed owing to the captive method, the later moulter is suspected to moult approximately 1 year after the earlier moulter. Earlier and later moulters were not different in growth of carapace width or chela height during the pubertal and terminal moult. Our moult and growth results of earlier and later moulters will be useful for predicting recruitment to legally fishable (>90 mm carapace width) or valuable (hard-shelled adult) populations of male snow crab in the Sea of Japan.     

Exploitation of floral signals by crab spiders (Thomisus spectabilis, Thomisidae)

Flowers exhibit characteristics through which they exploit thesensory biases of pollinating insects, and both signaler andreceiver benefit from this interaction, either through reproductiveservice or food reward. However, the preferences of pollinatorsfor certain flower traits such as color or odor might be exploitedby predators that target pollinating insects. Crab spiders,Thomisus spectabilis, position themselves on flowers to preyon pollinators such as honeybees, Apis mellifera. We gave bothhoneybees and crab spiders the choice between two randomly chosenwhite Chrysanthemum frutescens, including olfactory signalsin one experiment and excluding odor in a second experiment.When olfactory signals were included, crab spiders and honeybeesclearly preferred the same flower out of a pair. However, agreementlevel was at chance in the absence of olfactory signals. Wealso analyzed the visual flower characteristics that might influencethe decision of the animals. Neither the size of flowers (diameterof flower and diameter of reproductive flower center) nor thereflectance properties (receptor excitation values in ultraviolet,blue, and green; overall brightness) influenced the choicesof crab spiders and honeybees. Therefore, odor seems to be thefloral signal that bees use to identify high-quality flowersand that crab spiders exploit to encounter honeybees.     

Cichlids do not adjust reproductive skew to the availability of independent breeding options

Helpers in cooperatively breeding species forego all or partof their reproduction when remaining at home and assisting breedersto raise offspring. Different models of reproductive skew generatealternative predictions about the share of reproduction unrelatedsubordinates will get depending on the degree of ecologicalconstraints. Concession models predict a larger share when independentbreeding options are good, whereas restraint and tug-of-warmodels predict no effects on reproductive skew. We tested thesepredictions by determining the share of reproduction by unrelatedmale and female helpers in the Lake Tanganyika cichlid Neolamprologuspulcher depending on experimentally manipulated possibilitiesfor helper dispersal and independent breeding and dependingon helper size and sex. We created 32 breeding groups in thelaboratory, consisting of two breeders and two helpers each,where only the helpers had access to a nearby dispersal compartmentwith (treatment) or without (control) breeding substrate, usinga repeated measures design. We determined the paternity andmaternity of 1185 offspring from 47 broods using five to nineDNA microsatellite loci and found that: (1) helpers participatedin reproduction equally across the treatments, (2) large malehelpers were significantly more likely to reproduce than smallhelpers, and (3) male helpers engaged in significantly morereproduction than female helpers. Interestingly, in four broods,extragroup helper males had fertilized part of the brood. Nohelper evictions from the group after helper reproduction wereobserved. Our results suggest that tug-of-war models based oncompetition over reproduction within groups describe best thereproductive skew observed in our study system. Female breedersproduced larger clutches in the treatment compared to the controlsituation when the large helpers were males. This suggests thatmale breeder-male helper reproductive conflicts may be alleviatedby females producing larger clutches with helpers around.     

Resource partitioning between sexes in the "unconventional" hermit crab, Calcinus tubularis

Together with Calcinus verrilli from Bermuda, the Mediterraneanhermit crab C. tubularis is unique in that it exhibits a sharpsexual dimorphism in resource use in which one sex, the male,occupies loose shells and the other, the female, occupies attachedtubes. Field surveys and laboratory experiments addressed twoissues that help understand male-female resource partitioningin this species. First, the value of shells as shelters differsbetween sexes. Shells furnish mobility to their inhabitants,but also greater opportunities to grow because they can providelarger size than can the tubes, whereas both mobility and largedimensions are crucial selective factors for the reproductiveoutcomes of males only. In fact, egg production seemed not tobe affected by females' sedentary life and the number of eggswas not related to the female size. Second, previous shelterexperience plays a role in reducing male-female competition.I found that individual crabs, once presented simultaneouslywith a shell and a tube, more likely selected the type of shelterthat they were collected in. Then, in male-female competitionexperiments, pairs composed of one crab found in a shell andone crab found in a tube and offered a shell and a tube in conjunctionoccupied the housing in accordance to the shelter previouslyoccupied without any influence of their sex or size. The questionremains as to how, why, and when such a sexual dimorphism inthe distribution of C. tubularis between microhabitats did occurduring a hermit crab's life.