Does the handedness of the pebble crab Eriphia smithii influence its attack success on two dextral snail species?

The predaceous crab Eriphia smithii (Xanthidae) has one larger claw with molar teeth on either the right or the left cheliped, which it uses to crush the shell of prey. Whether the handedness of crabs affected successful predation on two snail species, Nerita albicilla (Neritidae) and Planaxis sulcatus (Planaxidae) was experimentally investigated. The fate of snails of each species was analysed by multiple logistic regression with three explanatory variables: handedness, shell-size index and individuality of crabs. No effect of handedness was detected in attacks on N. albicilla , probably as a result of the spherical and more symmetrical shell morphology of this species. In contrast, right-handedness contributed to greater attack success on P. sulcatus , which has more conical shells. Further investigation of how snail shells were broken revealed that left-handed crabs had more difficulty breaking the aperture of larger P. sulcatus , which was thought to cause the difference in attack success between right- and left-handed crabs. The advantages conferred by handedness are discussed.     

Age- and sex-associated variations in the directional asymmetry of rhesus macaque forelimb bones

Several investigators have questioned the significance of handedness as an explanation of directional forelimb asymmetries, yet little has been done to isolate other explanatory factors. In this investigation, we analyze 61 female and 76 male rhesus macaque skeletons for evidence of age- and/or sex-associated variations in ten forelimb bone measurements. All significant directional asymmetries are found to favor the right side. Although some of these asymmetries are found to favor the right side. Although some of these asymmetries are compatible with the interpretation of muscle hypertrophy associated with preferential use of the right forelimb, the overall pattern suggests that age- and sex-related ontogenetic factors deserve equal consideration. Significant sex differences in asymmetry means are present within and across age groups (juveniles, subadults, and adults), and numerous changes in asymmetry with age are also found. A pattern of decreasing asymmetry with age was found in males, with 40% of the ten measures being asymmetrical in juveniles, 30% in subadults, and 20% in adults. Among females, this pattern is reversed. No significant asymmetries were found for juvenile or subadult females, whereas 40% of the measures were asymmetrical in adult females. We conclude that greater consideration of age- and sex-related factors is necessary when drawing samples for the purpose of investigating asymmetries, and an awareness of trait-specific age and sex patterns of variation is necessary when citing forelimb asymmetries in demographically nonrepresentative populations as evidence of handedness or other behavioral asymmetries.     

Directional asymmetry in the forelimb of Macaca mulatta

Asymmetry was investigated in the forelimbs of 150 rhesus monkey (Macaca mulatta) skeletons using measurements of right and left humerii, radii, ulnae, second metacarpals, and femora. Seven of the ten forelimb dimensions were larger on the right than on the left side. Paired t-tests revealed that the mean of the right side was significantly larger than that for the left for two measurements of the ulna and two of the humerus. No measurement was significantly larger on the left than on the right side. These results indicate a small but significant asymmetry in the forelimb bones of rhesus monkeys and, as is the case for humans, the direction of asymmetry favors the right side. Our findings are consistent with an interpretation of hypertrophy of certain muscles and opens the question of whether rhesus monkeys preferentially use their right forelimbs for manipulative tasks that require manual dexterity, as is the case for humans. These forelimb skeletal asymmetries are discussed in light of the recent literature on cortical asymmetry and handedness in nonhuman primates.     

Maternal effect of low temperature on handedness determination in C. elegans embryos

C. elegans embryos, larvae, and adults exhibit several left-right asymmetries with an invariant dextral handedness, which first becomes evident in the embryo at the 6-cell stage. Reversed (sinistral) handedness was not observed among > 10,000 N2 adults reared at 16°C or 20°C under standard conditions. However, among the progeny of adults reproducing at 10°C, the frequency of animals with sinistral handedness was increased to ∼0.5%. Cold pulse experiments indicated that the critical period for this increase was in early oogenesis, several hours before the first appearance of left-right asymmetry in the embryo. Hermaphrodites reared at 10°C and mated with males reared at 20°C produced sinistral outcross as well as sinistral self-progeny, indicating that the low temperature effect on oocytes was sufficient to cause reversals. Increased frequency of reversal was also observed among animals developed from embryos lacking the egg shell. Possible mechanisms for the control of embryonic handedness are discussed in the context of these results, including the hypothesis that handedness could be dictated by the chirality of a gametic component. © 1996 Wiley-Liss, Inc.     

Relative growth,morphological sexual maturity,heterochely, and handedness in Panopeus occidentalis (Brachyura,Panopeidae)

Studies on relative growth and sexual maturity are important to understand the reproductive biology of a species. The aims of this study were to determine the relative growth and to estimate the size of Panopeus occidentalis at morphological sexual maturity, as well as to confirm whether this species demonstrates heterochely or handedness. Individuals were collected every two months from March 2013 to July 2014 in the intertidal estuarine zone of Cananeia, São Paulo, Brazil. The following measurements were taken: carapace width (CW), carapace length (CL), right and left cheliped propodus length (CPL), right and left cheliped propodus height (CPH), right and left cheliped propodus width (CPW), abdomen width (AW), and first pleopod length (FPL). The morphometric relationship used to estimate the size at morphological sexual maturity were CW vs. AW for females and CW vs. FPL for males; these comparisons yielded estimated CW values of 15.60 mm and 16.67 mm, respectively. Heterochely was observed but handedness was not present. The species has a major cheliped on one side, but the side is not constant. This study provides the first insights on the relative growth, sexual maturity, heterochely, and handedness on a population of P. occidentalis in a conserved area.     

Behavioral and morphological asymmetries in chukar Alectoris chukar copulation

Birds often exhibit greater reproductive tract development on the left side than right side. Behavioral evidence from the three species for which data has been published indicates that these species copulate more frequently on the left side of females than on the right side. Missing from the literature are studies that compare asymmetry in copulation behavior to asymmetry in reproductive tract morphology of the same individuals of both sexes within a single species. To better understand the potential for cryptic sexual selection to influence avian copulation, we measured asymmetries in chukar Alectoris chukar copulation using 24 male and 29 female chukar brought into captivity from the wild. Chukar copulated (n=37) more from the left side (n=30) of females than the right side (n=7). The left testis of males was consistently greater in size, mass and volume than the right testis. The left ovary and oviduct of females was consistently functional with no observable development of the right ovary or oviduct. Left-side bias in direction of copulation, larger left testes, and functional left vaginal openings may act in concert to deliver spermatozoa to the oviduct, promoting fertilization.     

Carcinophyly of fish of the genus Careproctus (Scorpaeniformes: Liparidae) in waters of southeastern Sakhalin (Sea of Okhotsk)

In waters of southeastern Sakhalin (Sea of Okhotsk), clutches of snailfishes of the genus Careproctus containing eggs at different stages of development or prolarvae were found on gills of the largest individuals (mainly in males) of three species of crabs (Lithodidae)-Paralomis verrilli (clutches were found in 1.4% of individuals), P. multispina (0.8%), and Lithodes covesi (11.6%). With an increase in the carapace width of lithodid crabs, the amount of eggs of snailfishes deposited in them and their diameter increase. More frequently, clutches are located from the left side of the carapace of crabs. Two-side clutches occur in larger individuals of crabs. No direct relationship between the presence of clutches of snailfishes on gills of lithodid crabs with necrosis or reduction of gills was revealed. The form of relations between lithodid crabs and Careproctus snailfishes was defined as “lodging” with different degree of impact on the host.     

The impact of limb autotomy on mate competition in blue crabs Callinectes sapidus Rathbun

Summary This study is the first to demonstrate experimentally that autotomy (self-amputation of a body part) adversely affects competition for mates. Experiments were conducted using blue crabs Callinectes sapidus Rathbun to examine the consequences of limb loss and pairing precedence on mate acquisition by males. Two adult males of equivalent size were introduced sequentially into pools containing a sexually-receptive female and observed after 24 h and 48 h. One male in each pair was left intact, while the other experienced: (1) no autotomy, (2) autotomy of one cheliped, or (3) autotomy of both chelipeds, one walking leg, and one swimming leg. In the absence of a competitor (first 24 h), both intact and injured males established precopulatory embraces with females. Intact males were highly successful (84–95%) in defending females from intact or injured intruders in the second 24 h period. Both autotomy treatments, however, significantly reduced the ability of males to defend females from intact intruders. Females in experiments suffered greater frequency of limb loss than did males. In the field, paired blue crabs showed significantly higher incidence of limb loss than unpaired crabs. Limb loss frequency increases with body size, and field observations indicated that larger males may be more successful than smaller males in obtaining females. Both experimental manipulations and field studies provide strong evidence for mate competition in this ecologically and commercially important portunid species.     

Population and reproductive biology of the fiddler crab Uca thayeri Rathbun, 1900 (Crustacea: Ocypodidae) in a tropical mangrove from Northeast Brazil

Population and reproductive biology of Uca thayeri Rathbun, 1900 were studied for the first time in a tropical mangrove. Absolute density, sex ratio, population structure, handedness, breeding season and fecundity were investigated. Seven transects were delimited in a mangrove area of the Pacoti River, Northeast of Brazil (3° 43′ 02″ S/38° 32′ 35″ W). On each transect, ten 0.25 m² squares were sampled on a monthly basis during low tide periods from September 2003 to August 2004. A total of 483 crabs were obtained, of which 250 were males, 219 non-ovigerous females, and 14 ovigerous females. The U. thayeri population presented bi-modal size frequency distribution, with males and non-ovigerous females not differing significantly size-wise. Ovigerous females were larger than males and non-ovigerous females. The overall sex ratio (1:1.07) did not differ significantly from the expected 1:1 proportion. The major cheliped was the right one in 50% of the males. The observed density was of 8.5 individuals/m², with the specimens being found mostly in shaded areas. Ovigerous females were found in 5 months of the year, coinciding with the rainy season, suggesting that the population of U. thayeri presents seasonal reproductive events. Juvenile crabs were more abundant during the dry period, while larger crabs were found mainly during the rainy period. The fecundity of the studied population was much smaller than that of subtropical populations of this species. The regression analysis shows that the number of eggs increases linearly with the increase of carapace width.     

Hair-trigger autotomy in porcelain crabs is a highly effective escape strategy

The benefits of autotomy, the voluntary shedding of limbs, havebeen adequately demonstrated in vertebrates but are poorlystudied in invertebrates. We provide strong experimental evidencefor an antipredatory benefit to autotomy in two porcelain crabs(Petrolisthes cinctipes and P. manimaculis). Since the outcomeof autotomy depends critically on the species of predator andprey involved, we first surveyed field populations of porcelaincrabs to identify ecologically relevant predators to use insubsequent experiments. We then examined the escape tacticsof the porcelain crabs in response to the four potential predatorswe identified, all larger predatory crabs. Cheliped autotomywas induced by three of the predator species (Cancer antennarius,Hemigrapsus nudus, Pachygrapsus crassipes); the fourth didnot attack porcelain crabs. Autotomy occurred in response to34% of all attacks, and in 67% of attacks in which the preywas held at some point by only the cheliped. Autotomy was ahighly effective escape mechanism against these predators;58 of 59 crabs that autotomized escaped, usually while thepredator stopped pursuit to eat the shed cheliped. Relianceon autotomy as a primary mechanism for escaping predators maybe particularly common in small crabs that cannot adequatelydefend themselves by other means and in suspension-feedingcrabs that do not need their chelipeds for feeding.     

Post-settlement effects of habitat type and predator size on cannibalism of glaucothoe and juveniles of red king crab Paralithodes camtschaticus

Postlarval (glaucothoe) and juvenile (first crab stage, C1) red king crab Paralithodes camtschaticus actively select structurally complex substrata for settlement. Such habitats may provide them with shelter from predation during critical early stages. We tested this hypothesis by placing glaucothoe and juvenile crab in aquaria with or without natural or artificial habitats, and with or without predators (1-3-year-old red king crab) of two different sizes. Predators caused increased mortality of glaucothoe, but predator size, habitat presence and habitat type had no effect on survival. Predators caused significant mortality of C1 crabs in the absence of habitat, and mortality was inversely related to predator size. Density of glaucothoe on habitats was similar with or without predators, but density of C1 crab on habitats was higher than that of glaucothoe, and increased in the presence of large predators. Active selection for complex substrata by settling glaucothoe does not reduce cannibalism, but may pre-position them for improved survival after metamorphosis. In contrast, juvenile crabs modify their behavior to achieve higher densities in refuge habitats, which tends to dampen the effect of predation. These survival strategies may have evolved to compensate for the greater risk of predation in open habitats.     

Persistence of subtle departures from symmetry over multiple molts in individual brachyuran crabs: Relevance to developmental stability

Fluctuating asymmetry (FA) is generally viewed as a population-level character. It is described by some measure of the variance of the difference between the right and left sides for a collection of individuals. Very little is known of the developmental origins of FA, despite the fact that FA is widely used to estimatedevelopmental stability. We present a novel technique for examining the growth trajectory of the asymmetries that give rise to FA, and we explore two sample data sets for the brachyuran crabHemigrapsus nudus. We have traced the fate of these small, random deviations from perfect symmetry through three successive molts of individual crabs. Invertebrates that molt, and hence grow in discrete steps, provide an easily preserved record of their growth. Model II regressions of measurements from one molt versus the previous molt can help describe the stability of subtle departures from symmetry over time. Although any number of different patterns may occur, we identify four general cases: a) asymmetries vary at random throughout growth (random determination), b) asymmetries remain unchanged in sign and magnitude (constant asymmetries), c) asymmetries increase in proportion to character size and hence increase with growth (size-dependent asymmetries), and d) asymmetries persist, but are reduced in magnitude (damped asymmetries). Data from tenHemigrapsus nudus, measured for between 21 and 28 metrical, limb-segment characters over three successive molts, yielded associations most similar to our pattern ‘b’, although some subtle departures in the direction of pattern ‘c’ were also observed. Persistent asymmetries accounted for 26% and 20% of the variance among asymmetries between molts 1 and 2, and molts 2 and 3 respectively. Thus, in spite of large and rapid increments in the external size of the crab, these subtle asymmetries tended to persist in both direction and magnitude, from molt to molt. This result suggests either i) that individual crabs have a genetic predisposition towards asymmetry in a particular direction but contribute to a continuous and normal distribution ofR-L differences at the population level, or ii) that these subtle asymmetries arose at some earlier ontogenetic stage and were preserved through growth. Either interpretation has important ramifications for the study of FA. The first suggests that under some circumstances FA may not provide a valid measure of developmental instability, because subtle departures from symmetry in an individual may have a genetic basis. The second implies that subtle departures from bilateral symmetry are not ‘corrected’ as an individual grows.     

A case of malformation in Pachycheles serratus (Decapoda: Porcellanidae)

An adult male of Pachycheles serratus with a malformation on the right cheliped was found during a collection of anomuran crabs in coastal waters of the peninsula de Macanao, Margarita island, Venezuela. The specimen was found at La Carmela beach (11 degrees 04'N-64 degrees 20'W), and featured a bifurcated fixed finger on the right cheliped.     

The development of handedness in left/right asymmetry

The development of handed asymmetry requires a special mechanism for consistently specifying a difference between left and right sides. This is to be distinguished from both random asymmetry, and from those left/right differences that are mirror symmetrical. We propose a model for the development of handedness in bilateral animals, comprising three components. (i) A process termed conversion, in which a molecular handedness is converted into handedness at the cellular level. A specific model for this process is put forward, based on cell polarity and transport of cellular constituents by a handed molecule. (ii) A mechanism for random generation of asymmetry, which could involve a reaction-diffusion process, so that the concentration of a molecule is higher on one side than the other. The handedness generated by conversion could consistently bias this mechanism to one side. (iii) A tissue-specific interpretation process which responds to the difference between the two sides, and results in the development of different structures on the left and right. There could be direct genetic control of the direction of handedness in this model, most probably through the conversion process. Experimental evidence for the model is considered, particularly the iv mutation in the mouse, which appears to result in loss-of-function in biasing, and so asymmetry is random. The model can explain the abnormal development of handedness observed in bisected embryos of some mammalian, amphibian and sub-vertebrate species. Spiral asymmetry, as seen in spiral cleavage and in ciliates, involves only conversion of molecular asymmetry to the cellular and multicellular level, with no separate interpretation step.     

Specification of left-right asymmetry in the embryonic gut of Drosophila

Most animals exhibit stable left-right asymmetries in their body. Although significant progress has been made in elucidating the mechanisms that set up these asymmetries in vertebrates, nothing is known about them in Drosophila. This is usually attributed to the fact that no reversals of stable left-right asymmetries have been observed in Drosophila, although relevant surveys have been carried out. We have focused on the asymmetry of the proventriculus in the embryonic gut of Drosophila, an aspect of left-right asymmetry that is extremely stable in wild-type flies. We show that this asymmetry can be reversed by mutations in the dicephalic and wunen genes, which also cause reversals in the antero-posterior axis of the embryo relative to its mother. This is the first observation to suggest that left-right asymmetries in Drosophila can be reversed by genetic/developmental manipulations. It also suggests that maternal signals may initiate the specification of some left-right asymmetries in the embryo.     

Chimpanzee (Pan troglodytes) Precentral Corticospinal System Asymmetry and Handedness: A Diffusion Magnetic Resonance Imaging Study

Background

Most humans are right handed, and most humans exhibit left-right asymmetries of the precentral corticospinal system. Recent studies indicate that chimpanzees also show a population-level right-handed bias, although it is less strong than in humans.

Methodology/Principal Findings

We used in vivo diffusion-weighted and T1-weighted magnetic resonance imaging (MRI) to study the relationship between the corticospinal tract (CST) and handedness in 36 adult female chimpanzees. Chimpanzees exhibited a hemispheric bias in fractional anisotropy (FA, left>right) and mean diffusivity (MD, right>left) of the CST, and the left CST was centered more posteriorly than the right. Handedness correlated with central sulcus depth, but not with FA or MD.

Conclusions/Significance

These anatomical results are qualitatively similar to those reported in humans, despite the differences in handedness. The existence of a left>right FA, right>left MD bias in the corticospinal tract that does not correlate with handedness, a result also reported in some human studies, suggests that at least some of the structural asymmetries of the corticospinal system are not exclusively related to laterality of hand preference.     

Common Variants in Left/Right Asymmetry Genes and Pathways Are Associated with Relative Hand Skill

Humans display structural and functional asymmetries in brain organization, strikingly with respect to language and handedness. The molecular basis of these asymmetries is unknown. We report a genome-wide association study meta-analysis for a quantitative measure of relative hand skill in individuals with dyslexia [reading disability (RD)] (n = 728). The most strongly associated variant, rs7182874 (P = 8.68×10⁻⁹), is located in PCSK6, further supporting an association we previously reported. We also confirmed the specificity of this association in individuals with RD; the same locus was not associated with relative hand skill in a general population cohort (n = 2,666). As PCSK6 is known to regulate NODAL in the development of left/right (LR) asymmetry in mice, we developed a novel approach to GWAS pathway analysis, using gene-set enrichment to test for an over-representation of highly associated variants within the orthologs of genes whose disruption in mice yields LR asymmetry phenotypes. Four out of 15 LR asymmetry phenotypes showed an over-representation (FDR≤5%). We replicated three of these phenotypes; situs inversus, heterotaxia, and double outlet right ventricle, in the general population cohort (FDR≤5%). Our findings lead us to propose that handedness is a polygenic trait controlled in part by the molecular mechanisms that establish LR body asymmetry early in development.     

Asymmetry of skeletal effects of Dominant hemimelia

BACKGROUND: Dominant hemimelia (Dh) is a dominant mutation that arose spontaneously in mice; Dh animals exhibit reduced numbers of lumbar vertebrae and preaxial hindlimb defects. Absence of spleen occurs in both Dh/+ and Dh/Dh animals. This study was undertaken to characterize asymmetry of skeletal defects in the Dh mouse, specifically hindlimb asymmetries in association with axial defects. METHODS: A total of 29 Dh/+ and 100 +/+ fetuses (gestational day [GD] 18) were identified by phenotype and linked DNA and their skeletons were analyzed. RESULTS: The results revealed an asymmetry of hindlimb skeletal defects in Dh/+ animals. In +/+ fetuses, the left and right tibia were symmetrical with 99.0% of the animals possessing 6 lumbar vertebrae. However, Dh/+ fetuses showed asymmetry in length of left and right tibia and a reduction to 5 lumbar vertebrae in 86.2% of animals. There was a range from mild to severe asymmetry as evidenced by direct comparison of the length of the left to the right tibia of each animal. Tibial shortening was greater on the left than the right in 65.5% of Dh/+ fetuses; only 20.7% had symmetrical tibia. Oligodactyly, defined as absence of the first or second toe, was similarly more frequent on the left. CONCLUSIONS: Asymmetry is characteristic of many human limb malformations, although analysis of the molecular basis is difficult. Therefore, Dh/+ mice, which exhibit reduced numbers of lumbar vertebrae, asymmetric hindlimb defects, and complete absence of spleen, provide an important model for studying the relationship between axial patterning and asymmetric skeletal defects.     

The timing of decisions during shell investigation by the hermit crab, Pagurus bernhardus

Factors which influence decisions by hermit crabs concerning whether to approach, investigate and enter another shell have been investigated by systematically varying the size of the shell in possession and the size of the shell being offered. The probability that a crab will approach or enter the shell depends on both variables. Investigation after contact, however, depends only on the size of the shell in possession. Durations of each stage of the sequence are negatively correlated with the numbers of crabs in each experimental group performing the next stage. For the final sequence of aperture investigatory activities, the fewer crabs entering the offered shell, the greater the number of investigatory acts performed by those crabs. In addition to variation in number of acts and duration of investigation, specific use of appendages varies according to the experimental situation. Thus naked crabs use the minor cheliped to investigate shells in preference to the major cheliped, which is normally used by housed crabs, with the major cheliped being reserved for defence by naked crabs. These data are discussed in terms of information collection and decisions made on the basis of that information.     

A Mathematical Modelling for the Cheliped Regeneration with Handedness in Fiddler Crab

An enormously developed giant cheliped with the other small one characterizes the adult male fiddler crab. Some experiments with artificial severances of cheliped indicate that such a handedness in the cheliped size is maintained even after the regeneration of severed cheliped. Other experimental researches give some results about an unknown physiological system which controls the emergence and the regeneration of the handedness in the cheliped size. In this paper, with two hypothesized factors relevant to the regeneration of a severed cheliped, we propose a simple mathematical model to describe the experimental result about the cheliped regeneration with a handedness after the cheliped severance for the fiddler crab. Our model gives a suggestion about an underlying system for the cheliped regeneration in the fiddler crab or some other crustacean species.