Frequency of moulting by shore crabs Carcinus maenas (L.) changes their colour and their success in mating and physiological performance

Male shore crabs, Carcinus maenas [Linnaeus, 1758], compete aggressively for access to receptive females to mate. Size is the single most important factor for the outcome of these conflicts, large males with carapace width (CW) over 60 mm being much more likely to gain access to receptive females than smaller males. To compete aggressively, large male shore crabs decrease moulting frequencies and may potentially terminate moulting to enter a state of anecdysis, in which further growth is suspended to increase reproductive output. This change from a “growth” strategy to a “reproduction” strategy results in the creation of two morphs, which can roughly be separated by their colouration. As the new exoskeleton created during moult is always green in appearance, crabs tend to be green in appearance during periods where they grow rapidly and moult frequently. Green crabs are found in all size classes. However, as the exoskeleton becomes older, the colour gradually changes to a darker red colour, and large crabs that have spent an extended period in intermoult are therefore often red in appearance. Also, the exoskeleton of red crabs exhibits a higher incidence of epibionts and wear and tear. Red crabs can also be found in all size classes, but their relative proportion in the population increases dramatically in size classes above 60-mm CW. Size for size, the red morph has a thicker carapace and larger master chelae than the green morph. Also, the reproductive indices (RI) for red crabs are higher than for green crabs, and they experience higher mating success. However, this mating success appears to be achieved at the expense of a lower physiological tolerance, green crabs being better adapted to deal with changes in the surrounding environment. This increased tolerance is not only observed with regard to natural variations in the habitats where shore crabs live, but green crabs also appear to be more tolerant to variations caused by anthropogenic pollution. Consequently, the shift from growth to reproduction exerts a profound effect on the behaviour, physiology and ecotoxicology of male shore crabs. The present paper reviews the studies conducted so far, proposes a mechanism by which some of these differences between the two morphs are created and discusses their ecological and ecotoxicological significance.     

Behavioral and electrophysiological experiments suggest that the antennular outer flagellum is the site of pheromone reception in the male helmet crab Telmessus cheiragonus

Sexually competent females of Telmessus cheiragonus (helmet crab) release two pheromones that elicit grasping and copulation behaviors in males (Kamio et al., 2000, 2002, 2003). Our study aimed to use behavioral and electrophysiological techniques to identify the site of reception of these sex pheromones. In behavioral experiments, either the inner or the outer flagella of the antennules were ablated bilaterally from male crabs, and responses of male crabs to female odor were examined. When the inner flagella were surgically ablated, the sexual response (i.e., grasping and copulation behavior) of male crabs was not significantly changed relative to control animals that had their second antennae ablated. In contrast, the sexual response was significantly reduced when the outer flagella of the antennules were ablated, suggesting that the outer flagellum is the receptor organ that detects the sex pheromones. In electrophysiological experiments, urine, which in females contains the pheromone that elicits grasping behavior by males but does not contain the pheromone eliciting copulation, whose release site is not known, was tested. Female and male urine as well as shrimp extract evoked phasic responses of chemosensory afferents innervating aesthetasc sensilla on the outer flagellum of male crabs. The response of the afferents had significantly higher magnitude and lower threshold when female urine was applied. Thus, behavioral and electrophysiological observations suggest that in male helmet crabs, the outer flagellum of the antennule is the chemosensory organ that detects female sex pheromone.     

Avoidance of post-coital cannibalism in the brachyurid paddle crab Ovalipes catharus

Sexual cannibalism was examined experimentally in the New Zealand paddle crab Ovalipes catharus, where it is the female that risks cannibalism during and after sexual activity. In this species copulation only occurs when the female is soft-shelled after moulting and she is most vulnerable to cannibalism. Male paddle crabs protect and copulate with the females until the females are no longer sexually receptive. Males appear able to identify a female with whom they have recently copulated. After a brief separation, significantly fewer females were cannibalized in cases where a female was returned to her original sexual partner than in those that had exchanged partners. The asymmetry, where females gain some protection during moulting but males receive no such advantage, may, at least partially, explain the skewed sex ratios which are sometimes found in samples of O. catharus from nature.     

Untersuchungen zur Häutungsphysiologie der dekapoden Krebse am Beispiel der StrandkrabbeCarcinus maenas

Under constant laboratory conditions, juvenile shore crabs moult at fixed intervals which depend upon their body size. During one moult every crab exhibits increases of the same relative amounts, independent of its absolute size. Basing on the predictable duration of the intermoult period, the morphological changes in the structure of the cuticle and the development of limb-buds, the intermoult period could be divided into 21 different stages. After studying the moulting rhythm in constant milieu, the influence of the following exogenous and endogenous factors upon the moulting rhythm and growth of normal and of eye-stalkless individuals was investigated: temperature, photoperiod, loss of pereiopods, feeding, and presence of larger specimens. From these investigations it became evident that the moulting rhythm is regulated by growth. The crabs are able to moult only after achieving a minimum of tissue growth. So long as this minimum growth is not achieved, a moult-inhibiting hormone is secreted and moulting is prevented. If the moult-inhibiting hormone is absent, moulting hormone is secreted and initiates a moult. Under dangerous conditions, the crabs are able to delay the next moult. Under unfavourable conditions they consume less food than normal. Therefore, the amount of tissue growth which is the necessary prerequisite for moulting is delayed, and continued release of moult-inhibiting hormone prevents the moult. Under conditions favourable for moulting, or demanding moult (e. g. after loss of many pereiopods) the crabs accelerate the moult. Temperature influences the moulting rhythm by indirect effects on the metabolic rate. During further investigations, the variation of the following parameters were determined quantitatively: content of moulting hormone in whole crabs; content of aminoacids, protein, glucose, Na⁺, K⁺, Mg⁺⁺ and Ca⁺⁺ in the hemolymph; pH and osmotic pressure in the hemolymph; and Ca⁺⁺ content in skeleton and whole crabs. All parameters mentioned — excepting pH and K⁺ content of the hemolymph — vary characteristically during the intermoult period. The titre of moulting hormone has 4 different maxima. Of all parameters, only the content of animoacids and protein in the hemolymph vary in the same way as the titre of the hormone. From these results the following conclusions are drawn: The moulting hormone not only initiates the moulting process, but controls it at several stages. Only protein metabolism seems to be under direct control of the moulting hormone which stimulates protein-synthesis. Chitin formation, regeneration, apolysis and ecdysis are indirectly controlled by the moulting hormone through protein metabolism. As in most of the other processes mentioned, the calcification of the new cuticle is not under the direct influence of the moulting hormone. The conclusion ofDigby (1966) that calcification in crabs is an electrochemical process, is confirmed.     

Regulation of sex-specific feeding behavior in fiddler crabs: physiological properties of chemoreceptor neurons in claws and legs of males and females

This study examined properties of chemoreceptor neurons in the claws and legs of the fiddler crabs Uca pugilator and U. pugnax. The primary goal was to establish the neural basis of previously observed greater female sensitivity to feeding stimulants, and secondarily to compare physiological properties of chemoreceptor neurons in these semi-terrestrial crustaceans with those of fully aquatic forms. Sensitivity of chemoreceptor neurons in claws and legs is sex-specific; individual neurons of females respond to lower stimulus concentrations than male chemoreceptor neurons, and equivalent concentrations elicit greater spiking in female vs male chemoreceptor neurons. Thus, the population of chemoreceptor neurons in females expresses lower thresholds and greater average sensitivity than in males. Greater sensitivity of claw neurons explains observations indicating that females continue to feed at food levels too low to stimulate males. Sensitivity differences in leg neurons of males vs females have no clear behavioral correlate, but suggest that females can orient to more dilute stimuli than males. Chemoreceptor neurons of fiddler crabs have low sensitivities and slow rates of adaptation compared to other crustaceans. Also, neurons in claws adapt less slowly than neurons in legs, which may reflect subtle differences in the chemical stimulus environment experienced by claws vs legs.     

Tuning breadth and sex-specific sensitivity in chemosensory neurons of male and female Uca pugnax

Chemosensory neurons of female fiddler crabs (genus Uca) display greater sensitivity to mixtures of food-related stimuli than do neurons in males. This phenomenon represents an interesting contrast to other sex-specific systems, which tend to be in response to cues associated with mating and parental care. This study examined the responses of chemosensory neurons in males and females to ten individual stimuli to determine if sex-specific responses were restricted to a few key compounds, or if the heightened sensitivity of females was broadly distributed. Neurons in males and females responded well to all stimuli, and although fiddler crabs are primarily herbivorous, highly efficacious physiological stimulants included amino acids and amines as well as carbohydrates most closely associated with plant material. The chemosensory neurons are characterized by broad tuning and relatively high response thresholds, when compared to other crustaceans. Most importantly, the investigations revealed a robust pattern in which female neurons displayed elevated responses to all stimuli. Tuning breadth was not shown to be sex-specific, nor were there detectable differences in over-all response profiles. The most likely explanation for these patterns is that the broad sex-specificity in Uca is produced via fundamental alterations in cellular properties associated with chemosensory transduction. Accepted: 29 June 1999     

Identification of sex-linked DNA markers by AFLP in Portunus trituberculatus

To investigate the mechanism of sex determination of crabs, amplified fragment-length polymorphism DNA analysis was applied to identify sex-specific markers for Portunus trituberculatus. Eleven pairs of primers were used to amplify DNA isolated from male and female crabs. A total of 481 and 499 fragments were amplified for females and males, respectively, and the ratios of polymorphic fragments were 20.59–65.52% and 27.27–62.22%, respectively. Distribution difference values between female and male groups for 10 polymorphisms were greater than 40%; Cluster analysis revealed that Nei's genetic distances for 57 polymorphisms clustered according to sex. The results indicate that a difference at the DNA level exists between female and male crabs and provide data for further study on sex determination.     

Sex-specific, counteracting responses to inbreeding in a bird

Inbreeding often depresses offspring fitness. Because females invest more than males in a reproductive event, inbreeding is expected to be more costly to mothers than fathers, creating a divergence between the reproductive interests of each sex and promoting sex-specific inbreeding strategies. Males and females may bias the probability of inbreeding by selecting copulation partners, and, in sexually promiscuous species, through male strategic sperm investment in different females and female selection of the sperm of different males. However, these processes are often difficult to study, and the way that different male and female strategies interact to determine inbreeding remains poorly understood. Here we demonstrate sex-specific, counteracting responses to inbreeding in the promiscuous red junglefowl, Gallus gallus. First, a male was just as likely to copulate with his full-sib sister as with an unrelated female. In addition, males displayed a tendency to: (i) initiate copulation faster when exposed to an unrelated female than when exposed to a sister, and (ii) inseminate more sperm into sisters than into unrelated females. Second, females retained fewer sperm following insemination by brothers, thus reducing the risk of inbreeding and counteracting male inbreeding strategies.     

Mating Effect on Sex Pheromone Production of the Oriental tobacco budworm,Helicoverpa assulta

This study was undertaken to clarify the suppression phenomenon of sex pheromone production after mating and its relationship to the physiological mechanism in adult females of Helicoverpa assulta, and determine the mating factor from males causing depletion of sex pheromonc production. Sex pheromone production of H. assulta females was mostly terminated in 3 hours after mating. Mated females maintained with a low titer of sex pheromone until 3 days when it started to increase again, which showed a characteristic of species mating more than once. The mated female again produced pheromone upon injection of pheromone biosynthesis activating neuropeptide (PBAN) or extracts of brain-suboesophageal ganglion complexes (Br-Sg) of mated female, which were shown similar pheromonotropic activities as compared with virgin females. These results indicated that the mating did not inhibit the receptivity of pheromone gland itself and PBAN biosynthesis in suboesophageal ganglion of the mated females. And it seems to support that the depletion of sex pheromone production is responsible for blocking of PBAN release from head. To investigate the mating factor from adult males, when extracts of reproductive organs of male were injected into hemocoel of virgin females evoking depletion of sex pheromone production as shown in mated female. The results suggest that a chemical substance(s) from the male reproductive organs could be responsible for the loss of sex pheromone biosynthesis in H. assulta.     

Male Aggression,Female Mimicry and Female Choice in the Rove Beetle,Aleochara curtula (Coleoptera,Staphylinidae)

Causes, consequences and pheromonal regulation of male contest and female choice in the staphylinid beetle, Aleochara curtula (Goeze), have been investigated in the field and in the laboratory. At the feeding and mating site (carcass) the sex ratio is male biased. Polyandry is affected by prolonged copulations, spermatophore plugs and anti-aphrodisiac pheromones transferred from the male, and by female repulse behaviour as well. Aggression of competing males leads to expulsion of inferior males from the carcass. Young, starved and multiply mated males, which need access to the food resource, produce the female sex pheromone. They release homosexual responses, but also avoid intrasexual aggression. On the other hand, females behave aggressively towards individuals bearing the female sex pheromone or repulse their copulatory attempts. Those males of insufficient physiological condition produce a lighter spermatophore and fertilize less eggs. The adaptive significance of female mimicry, male mating tactics, and female choice is discussed.     

The influence of temperature,sex and chela size in the foraging strategy of the shore crab,Carcinus maenas (L.)

Foraging rate was highly variable among shore crabs of the same size category and for individual crabs from day to day. Possible physiological reasons for this variability are discussed. Shore crab foraging rate, both in terms of mussels eaten per day and energy intake per day, was estimated to be higher at 17°C than at 10°C. The shape of diet curves and their mode for male shore crabs at 17°C closely resembled those for 10°C, indicating that the temperature increase had no effect on their previously demonstrated optimal foraging strategy.

Female and certain male shore crabs showed a preference for prey smaller than for other equivalent sized males. These suboptimally feeding male and female crabs attained a relatively higher prédation rate (mussels day^‐1), although their energy intake (KJ day^‐1) remained lower than that of optimally feeding males. Preferred mussel size, number of mussels eaten per day and energy intake were strongly related to master chela height. The diet curves for female and suboptimally feeding male shore crabs could be explained by these crabs’ proportionately smaller master chelae.     

Mating opportunity and the evolution of sex-specific mortality rates in a butterfly

Life history theory predicts that organisms should only invest resources into intrinsic components of life span to the degree that it pays off in terms of reproductive success. Here, we investigate if the temporal distribution of mating opportunities may have influenced the evolution of intrinsic mortality rates in the butterfly Pararge aegeria (Satyrinae). In this species, females mate only once and the frequency of male mating opportunities depends on the temporal emergence pattern of virgin females. As expected, in a population from Madeira where females emerge continuously throughout the year, there was no sex difference in adult life span, while in a Swedish population with synchronised female emergence, males had significantly shorter life spans compared to females. A logistic mortality model provided the best fit to the observed change in age-specific mortality and all categories reached an asymptotic mortality rate of a similar magnitude. However, the Swedish males reached this mortality plateau more rapidly than the other categories. External mortality, due to water and food limitation, affected the pattern of sex-specific mortality but males from Sweden still had higher rates of mortality compared to all other categories. We argue that selection on male longevity is likely to be weaker in Sweden because under synchronised emergence, all females emerge and mate within a short period of time, after which male reproductive value will quickly approach zero. On Madeira, however, male reproductive value decrease more slowly with age since the probability of finding a receptive female is constant over the year. Received: 29 July 1999 / Accepted: 23 August 1999     

Reproductive biology of the shore crab Carcinus maenas (Decapoda,Portunidae): a macroscopic and histological view

A histological study of the reproductive cycle of male and female shore crabs, Carcinus maenas (Linnaeus), was performed monthly on the South West coast of Ireland (from December 2006 to July 2008). The calculated sex ratio deviated from equality, 1:0.53, revealing a strong male bias. A system was devised, based on screening of tissue sections, to describe and stage gametogenic development. Histological examinations revealed that ovarian development occurred biannually, with a primary winter cycle in which the larger crabs reproduced and a secondary summer cycle, when smaller crabs reproduced. An association was observed where more of the larger specimens were caught in the summer months and the smaller specimens in the winter months, which inversely correlated with the segregated breeding cycles. There was strong evidence that mature male crabs could potentially copulate year round since all mature specimens, collected throughout the year, contained viable spermatozoa. Developmental stages of oogenesis and spermatogenesis were described to develop a practical gonadal index for this portunid crab, providing information on the biology of this species, which will be of benefit for fisheries management.     

Reproductive performance of male and female phenotypes in three sex chromosomal genotypes(XX, XY, YY) in the killifish, Oryzias latipes.

The present study investigates the juvenile and adult reproductive performance of Japanese killifish, Oryzias latipes, which were successfully sex-reversed by feeding them male or female sex hormones during the fry stage. Sexual maturation of these laboratory grown fish of three known genotypes (XX, SY, YY) occurred earliest in untreated genotypes, next in genotypes treated with male hormone (methyl testosterone) and female hormone (estrone), respectively. The delay in sexual maturation caused by early exogenous, sex-hormone treatment may represent a disturbance in the delicate neural-gonadal axis. However, the degree of response was also strongly influenced by the animal's sex chromosomal genotype. XX fish, normally phenotypic females, were affected more by the male hormone than by the female hormone. XY and YY fish, normally phenotypic males, were delayed more by the female hormone than by the male hormone. This specific sex hormone-sex genotype interaction also influenced adult reproductive performance (sperm or egg production). Differences in the metabolism of male and female sex hormones by the XX, XY, and YY fish are probably responsible for these interesting findings.     

Population features of Microphrys bicornutus Latreille, 1825 (Brachyura, Majidae) from Isla Margarita, Venezuela

The aim of this study is to characterize the population biology (sex ratio, size structure, relative growth of reproductive characters and realised fecundity) of the crab Microphrys bicornutusfrom Isla Margarita, Venezuela. Nearly 100 crabs were collected and fixed monthly during 1998. In the laboratory, crabs were sexed and measured: maximum carapace length, chelae length, chelae and abdomen width for females and first pleopod length for males. Additionally, 96 ovigerous females were collected to determine fecundity. The following measurements were taken for each female: body wet weight (BWW), body dry weight (BDW), egg wet weight (EWW), egg dry weight (EDW) and number of eggs (NE). The following determinations were made: relative fecundity (RF) = NE/BDW and reproductive output (RO) = (EDW/ BDW) × 100. Relative size at the onset of maturity (RSOM) was calculated as minimum ovigerous female/female's maximum size. The results indicate that the sex ratio is nearly 1:1 during the entire year. Ovigerous females and moulted males and females were observed throughout the year. Size frequency distribution and size at the onset of sexual maturity, estimated by relative growth of the sexual secondary characters and RSOM value, differed from those estimated in studies done in Jamaica and Buchuaco, Venezuela.     

Ecdysteroids as a component of the genital sex pheromone in two species of hard ticks,Dermacentor variabilis andDermacentor andersoni (Acari: Ixodidae)

Neutering of part-fed females eliminated copulatory behavior inDermacentor variabilis (Say) andD. andersoni Stiles males. Extracts from the anterior reproductive tracts of part-fed (7 days) females partly restored the male copulatory behavior in conspecific neutered females. Similar extracts from unfed females did not restore the behavior, suggesting that the pheromone was produced during feeding. Perception of the genital sex pheromone by sensillae on the male cheliceral digits was confirmed by electrophysiological techniques.Males ofD. variabilis andD. andersoni responsed positively to authentic ecdysone and 20-hydroxyccdysone (20HE) in neutered female bioassays. Responses to sterols were significantly lower than to ecdysteroids. Electrophysiological assays suggest a sensitivity of males to high doses of ecdysteroids. The strongest responses were to 20HE in both species. Ecdysteroids, specifically ecdysone and 20HE were shown to be present in the anterior reproductive tracts in excess of amounts that could be explained by mere hemolymph contamination. Ecdysteroids were also found in washings of the vaginal lumen of these two species.Dermacentor andersoni females contained larger amounts of ecdysteroids thanD. variabilis females. 20-hydroxyecdysone and possibly ecdysone appear to be components of the genital sex pheromone ofD. variabilis andD. andersoni. Species recognition may be facilitated by these components, but the complete mechanism is not yet fully understood.Supported by grant AI10, 986 from the National Institute of Allergy and Infectious Diseases, National Institutes of Health, U.S. Public Health Service, Department of Health and Human Services, Bethesda, Maryland.     

赤腹松鼠血清性激素水平的季节差异

正季节性繁殖是动物适应环境的具体表现,也是动物维持其种群发展的重要策略。生殖激素和动物的繁殖活动关系密切,协调实现繁殖后代的机能(赖平等,2012),如配子的发生、成熟与排出及受精、妊娠、分娩与泌乳等性行为活动。例如雄性甘肃鼢鼠(Myospalax cansus)血清中睾酮(程志兴,2009)和黄山短尾猴(Macaca thibetana)血清中孕酮含量(夏东坡,2007)在交配期间显著升高,     

The antagonism between somatic growth and testicular activity during different phases in intermoult (stage C4) in sexually mature freshwater crab,Paratelphusa hydrodromous

Summary

In adult males of the freshwater crab Paratelphusa hydrodromous, judging from testicular activity, November-May is the reproductively inactive season and June/July through October the active season. The reproductively active period also represents the mating season of this species. In males over one year old, moulting is an annual event scheduled during June/July and the individual remains in intermoult from July/August to May. This intermoult comprises two phases: (1) the reproductive phase (July/August through October) when the physiological emphasis is more on reproduction and much less on somatic growth (as evidenced by the ability to regenerate autotomized walking limb), (2) the somatic phase (February/March through May) when the emphasis is more on somatic growth and much less on reproduction. Thus there exists an antagonism between reproduction and somatic grown in male P. hydrodromous as already established in the female. Experimental studies suggest that the testicular inactive phase of P. hydrodromous is caused by a possible increase of titre of the gonad-inhibiting hormone (GIH) along with a decrease in titres of the gonad-stimulating hormone (GSH) and androgenic hormone (AH).     

A physiological marker for quantifying differential reproductive investment between the sexes in Yellow-legged gulls (Larus michahellis)

Asymmetry between males and females in the energy they invest initially in reproduction has resulted in the evolution of differing reproductive strategies (caring females vs. competitive males). However, parental care in many birds is shared by both sexes suggesting that male energy expenditure in agonistic behaviors and courtship feeding might compensate female costs of clutch production. Here, we tested the hypothesis that initial investment in reproduction by both sexes in the Yellow-legged Gull (Larus michahellis), a species with biparental care, is similar from a physiological perspective. In this income breeder, female and male reproductive investment during early breeding can be ultimately related to muscular activity (local foraging effort required for clutch production in females and courtship feeding and agonistic behaviors in the case of males). Thus, we evaluated sex-specific patterns of creatine kinase (CK, IU/L) levels in plasma, an indicator of physical effort associated with muscular activity dependent behaviors, through incubation as a reflection of the physiological response of both sexes to the reproductive investment they made up to clutch completion. Raw levels of CK were related to plasma levels of total proteins (TP, g/dL) to account for the differential physiological state of individuals when sampled (i.e. differential dehydratation). Female costs of clutch production were associated with post-laying levels of CK/TP. We grouped females according to their relative investment in clutch production: < 15.8%, 15.8 to 17.3% and > 17.3% of field metabolic rate; which showed increasing values of CK/TP (24.6, 53.1 and 66.0 IU/g, respectively). Moreover, we found similar CK/TP trends throughout incubation for both sexes (CK/TP = 50.2− [3.3 × days from laying]) suggesting similar physiological responses to reproductive effort and, therefore, analogous sex-specific initial investment. Thus, male investment in agonistic behaviors and courtship feeding apparently equaled female investment in clutch production. The use of CK measurements is revealed as a useful approach to investigating overall investment in reproduction for both sexes, providing further insights into our comprehension of reproductive strategies in seabirds.