Morphology and muscle stress of chelae of temperate and tropical stone crabs Menippe mercenaria

Chela morphology and muscle stress were compared between temperate and tropical populations of stone crabs Menippe mercenuriu (Say) to test whether environmental differences might result in greater crushing strength in the tropics. Such differences include increased crab diversity in the tropics (which might lead to greater fighting among congeners), increased prey exoskeleton calcification in the tropics, and year round chela use in the tropics as opposed to seasonal chela use in the temperate population.
No latitudinal differences were found in any aspect of chela morphology, including relative chela size, mechanical advantage, apodeme surface area, and angle of muscle fibre pinnation. Summer measurements of crusher chela muscle stress were also similar between the two populations.
The maximum muscle stress determined for M. mercenariu was 220 N ∼r n -∼, much higher than stress levels previously reported for crustaceans. Other researchers have typically measured forces either from autotomized chelae or by measuring forces required to open a closed chelae. I have determined muscle stress using a force transducer that measures active gripping strength in live crabs.     

Structure and function of the chelae and chela closer muscles of the shore crab Carcinus maenas (Crustacea: Brachyura)

Major and minor chelae of Carcinus maenas were investigated with respect to mechanical advantage, angles of pennation and sarcomere lengths of the closer muscle fibres, maximum pinching forces induced by nervous stimulation and maximum stress in the closer muscles. Major chelae differed from minor chelae in all these factors. Mean mechanical advantages of major and minor chelae were 0·36 and 0·26, respectively. Angles of pennation were greater at the distal ends of closer muscles and declined proximally: in half open major and minor chelae distal to proximal angles were about 35–26° and 29–19°, respectively. Mean sarcomere lengths in closer muscles of fully closed major and minor chelae were significantly different at 7·6 and 6·0 fim, respectively; there was great variation within chelae and between crabs. Major chelae pinched more strongly than minor chelae due to higher mechanical advantages, larger closer apodemes and greater stress in the muscle. Mean maximum stresses in major and minor chelae were 677 and 474 kN/m², respectively. The biggest stress recorded–1057 kN rrr²–was in a major chela. Pinching forces induced by injecting a high K⁺ and caffeine ringer were about 20% weaker than nervously stimulated pinches.     

Leverage and muscle type in crab chelae (Crustacea: Brachyura)

The chelae of Cancer pagurus and Macropipus depurator were examined with respect to mechanical advantage. The closer muscles were investigated with respect to sarcomere length in the constituent fibres and to the force developed by the whole muscle during isometric contraction. Cancer chelae have a relatively high mechanical advantage, 0.329 ± 001. Cancer closer muscles contain a high proportion of fibres with long sarcomeres, mean lengths mostly falling between 12 and 15 μm, and develop a maximum stress of about 496 kN.m⁻² during contraction. These figures are typical for "slow" crustacean muscle. The chelae of M. depurator are dimorphic. In one, the strong chela, the mechanical advantage is 0.248 ± 0.066 while in the other, the fast chela, the mechanical advantage is 0.177 ± 0.006. M. depurator closer muscles contain fibres with mean sarcomere lengths mostly falling between 6 and 10 μm. The muscle develops a maximum stress of about 145 kN.m⁻² during contraction. These figures are typical of "intermediate" crustacean muscles. "Fast" muscle fibres with short sarcomeres (about 30 um) were found in the chelae of both Cancer and M. depurator but were much commoner in the latter. Thus in Cancer a high mechanical advantage is correlated with slow muscle while in M. depurator lower mechanical advantages are broadly correlated with faster muscle. Consistent correlation between mechanical advantage and muscle type in the dimorphic chelae of M. depurator , however, is lacking. No consistent regionation of fibres with similar properties was found in the muscles.     

Size,shape, and sex‐dependent variation in force production by crayfish chelae

The ability to generate large closing forces is important for many animals. Several studies have demonstrated that bite or pinching force capacity is usually related to the linear dimensions of the closing apparatus. However, relatively few studies have applied geometric morphometrics to examine the effects of size‐independent shape on force production, particularly in studies of crustacean pinching force. In this study, we utilized traditional and geometric morphometric techniques to compare the pinching force of Procambarus clarkii crayfish to their chela morphology. We found that males possessed larger chelae and pinched harder than females, but that their chela shape and size were weak predictors of strength. Female pinching force was significantly affected by both chela size and shape, with shape variation along the short axis of the claw contributing most to pinching force. We discuss our results in the context of reliable signaling of strength by males and females, and the different selective forces acting on chela shape in the two sexes.     

A study of normal and regenerative growth in the pistol-crab,alpheus dentipes (Guèr.), with special reference to the phenomenon of chela reversal

Summary An analysis has been made of the normal growth of the thoracic appendages in the pistol-crab,Alpheus dentipes. The chelae are the only asymmetrical appendages and, within an extensive size range, the growth-coefficient of the large chela (crusher) is smaller than that of the small chela (nipper), even in the most rapidly growing dimensions of the segment containing the growth-centre.The partition coefficient for growth in certain dimensions of the chelae is not constant but is reduced progressively during the growthperiod, due (it is suggested) to the impracticability of maintaining the intensive heterogony developed during the early stages of growth.Regenerative growth of the chelae has been studied quantitatively after autotomy of one, the other, or both normal chelae, effected once only or repeatedly as new chelae appeared at moulting. New chelae are regenerated repeatedly with little or no waning of growth intensity but in continuously growing chelae, growth intensity is substantially reduced at the end of the instar of autotomy. The amount of growth is not correlated with thenumber of days available for regeneration. When both chelae regenerate simultaneously the larger one grows relatively slower than the smaller one to the end of the instar of autotomy but afterwards this condition is reversed. When the nipper alone is autotomised, growth of the crusher is inhibited and shrinkage may occur. Crusher autotomy, on the other hand, stimulates nipper growth and to an extent which varies with the dimension.The problem of chela reversal has been discussed and certain suggestions have been preferred.     

Choose Your Weapon: Defensive Behavior Is Associated with Morphology and Performance in Scorpions

Morphology can be adaptive through its effect on performance of an organism. The effect of performance may, however, be modulated by behavior; an organism may choose a behavioral option that does not fully utilize its maximum performance. Behavior may therefore be decoupled from morphology and performance. To gain insight into the relationships between these levels of organization, we combined morphological data on defensive structures with measures of defensive performance, and their utilization in defensive behavior. Scorpion species show significant variation in the morphology and performance of their main defensive structures; their chelae (pincers) and the metasoma (“tail”) carrying the stinger. Our data show that size-corrected pinch force varies to almost two orders of magnitude among species, and is correlated with chela morphology. Chela and metasoma morphology are also correlated to the LD50 of the venom, corroborating the anecdotal rule that dangerously venomous scorpions can be recognized by their chelae and metasoma. Analyses of phylogenetic independent contrasts show that correlations between several aspects of chela and metasoma morphology, performance and behavior are present. These correlations suggest co-evolution of behavior with morphology and performance. Path analysis found a performance variable (pinch force) to partially mediate the relationship between morphology (chela aspect ratio) and behavior (defensive stinger usage). We also found a correlation between two aspects of morphology: pincer finger length correlates with the relative “thickness” (aspect ratio) of the metasoma. This suggests scorpions show a trade-off between their two main weapon complexes: the metasoma carrying the stinger, and the pedipalps carrying the chelae.     

Dishonest signals of strength in male slender crayfish (Cherax dispar) during agonistic encounters

Many animals resolve disputes without combat by displaying signals of potential strength during threatening displays. Presumably, competitors use each other's displays to assess their relative strengths, and current theory predicts that these signals of strength should generally be honest. We tested this prediction by investigating the relationships among morphology, performance, and social dominance in males of the slender crayfish Cherax dispar. Crayfish routinely use their enlarged front claws (chelae) for both intimidation and fighting, making this species ideal for studying the honesty of weapon size. We evaluated five competing models relating morphological and physiological traits to dominance during paired competitive bouts. Based on the best model, larger chelae clearly resulted in greater dominance; however, chela strength had no bearing on dominance. Thus, displays of chela size were dishonest signals of strength, and the enlarged chelae of males seemingly function more for intimidation than for fighting. In addition, an analysis of the performance of isolated chela muscle showed that muscle from male crayfish produced only half the force that muscle from female crayfish produced (236.6+/-26.4 vs. 459.5+/-71.6 kN m(-2)), suggesting that males invest more in developing larger chelae than they do in producing high-quality chela muscle. From our studies of crayfish, we believe dishonest signaling could play a greater role in territorial disputes than previously imagined.     

Decapod crustacean chelipeds: an overview

The structure, growth, differentiation and function of crustacean chelipeds are reviewed. In many decapod crustaceans growth of chelae is isometric with allometry level reaching unity till the puberty moult. Afterwards the same trend continues in females, while in males there is a marked spurt in the level of allometry accompanied by a sudden increase in the relative size of chelae. Subsequently they are differentiated morphologically into crusher and cutter making them heterochelous and sexually dimorphic. Of the two, the major chela is used during agonistic encounters while the minor is used for prey capture and grooming. Various biotic and abiotic factors exert a negative effect on cheliped growth. The dimorphic growth pattern of chelae can be adversely affected by factors such as parasitic infection and substrate conditions. Display patterns of chelipeds have an important role in agonistic and aggressive interactions. Of the five pairs of pereiopods, the chelae are versatile organs of offence and defence which also make them the most vulnerable for autotomy. Regeneration of the autotomized chelipeds imposes an additional energy demand called “regeneration load” on the incumbent, altering energy allocation for somatic and/or reproductive processes. Partial withdrawal of chelae leading to incomplete exuviation is reported for the first time in the laboratory and field inMacrobrachiumspecies.     

Crabs grab strongly depending on mechanical advantages of pinching and disarticulation of chela

A small morphological variation of an organ may cause a major change of its function in animal evolution. The function of decapod chela varies considerably among taxa, between sex, and even within an individual, but also retains a simple mechanism of motion. Therefore, the decapod chela is a suitable structure to study the evolutionary process of functional diversifications, although the relationship of form and function is inadequately understood, yet. We estimated the mechanical advantages of pinching and passive disarticulation resistance, and chela size relative to the carapace in 317 chelae of 168 decapod specimens, and compared these indices with the functions of each chela. Our study revealed that mechanical advantages of pinching efficiency and passive disarticulation resistance were greatest in shell‐crushing chelae, followed by gripping and pinching chelae, whereas the chela size relative to the carapace was not related to differences among these functions. We also found that the chelae are designed to retain the ratio between depth and width of the proximal dactylus. In the evolutionary process of decapods, the diversifications of chela functions were accompanied by the diversifications of the mechanical advantages, and played an essential role in their ecological diversification. J. Morphol. 277:1259–1272, 2016. © 2016 Wiley Periodicals, Inc.     

Heterochely and handedness in the shore crab Carcinus maenas (L.) (Crustacea: Brachyura)

A large sample ofadult male Carcinus maenas was 79% right-handed and 21% left-handed. A separate sample of 207 intact adult males was divided into left-handed and right-handed crabs and four measurements were taken from all major and minor chelae. Correlation and regression analyses against carapace width on log-transformed data showed that major chelae of right-handed crabs grow proportionately higher with increasing size and the ideal mechanical advantage increases; concurrently, the fingers of the minor chelae grow proportionately longer. The data for left-handed crabs showed greater variability, especially for minor chelae, providing evidence for the concept that left-handedness arises by reversal of handedness following loss of the major chela from the right-hand side. Records of handedness in large samples of non-ocypodid heterochelous brachyuran crabs show a preponderance of right-handedness.     

Claw asymmetry in lobsters: case study in developmental neuroethology.

An enduring debate in the study of development is the relative contribution of genetic and epigenetic factors in the genesis of an organism, that is, the nature vs. nurture debate. The behavior of the paired claws in the lobster offers promising material for pursuing this debate because of the way they develop. The paired claws and their closer muscles are initially symmetrical; both are slender in appearance and have a mixture of fast and slow fibers in their closer muscles. During a critical period of development, they become determined into a major (crusher) and minor (cutter) claw and during subsequent development acquire their final form and behavior: The crusher becomes a stout, molar-toothed claw capable of closing only slowly because its closer muscle has 100% slow fibers while the cutter becomes a slender, incisor-toothed claw capable of closing rapidly because its closer muscle has 90% fast fibers. Our initial hypothesis was that the more active claw became the crusher and its less active counterpart the cutter. Presumably, nerve activity would influence muscle transformation, which in turn would influence the exoskeleton to which they attach and hence claw morphology. Curtailing nerve activity to the claw prevented crusher development, while reflex activation of a claw promoted its development; both results support the notion that nerve activity directly regulates claw form and function. This is not, however, the case, for when both claws were reflexly exercised neither formed a crusher, signifying rather that bilateral differences in predominantly mechanoreceptive input to the paired claws somehow lateralized the claw ganglion [central nervous system (CNS)] into a crusher and cutter side. The side experiencing the greater activity becomes the crusher side while the contralateral side becomes the cutter and is also inhibited from ever becoming a crusher. This initial lateralization in the CNS is expressed, via as yet unknown pathways, at the periphery in claw morphology, muscle composition, and behavior. The critical period defines a time when the CNS is susceptible to being lateralized into a crusher and cutter side. Such lateralization is dependent upon experience of the environment in the form of mechanoreceptive input. In the absence of such experience, the CNS is not lateralized and paired cutter claws develop. Thus, while the critical period for crusher determination is genetically determined the actual trigger is influenced by experience.     

Claw asymmetry in lobsters: Case study in developmental neuroethology

An enduring debate in the study of development is the relative contribution of genetic and epigenetic factors in the genesis of an organism, that is, the nature vs. nurture debate. The behavior of the paired claws in the lobster offers promising material for pursuing this debate because of the way they develop. The paired claws and their closer muscles are initially symmetrical; both are slender in appearance and have a mixture of fast and slow fibers in their closer muscles. During a critical period of development, they become determined into a major (crusher) and minor (cutter) claw and during subsequent development acquire their final form and behavior: The crusher becomes a stout, molar-toothed claw capable of closing only slowly because its closer muscle has 100% slow fibers while the cutter becomes a slender, incisor-toothed claw capable of closing rapidly because its closer muscle has 90% fast fibers. Our initial hypothesis was that the more active claw became the crusher and its less active counterpart the cutter. Presumably, nerve activity would influence muscle transformation, which in turn would influence the exoskeleton to which they attach and hence claw morphology. Curtailing nerve activity to the claw prevented crusher development, while reflex activation of a claw promoted its development; both results support the notion that nerve activity directly regulates claw form and function. This is not, however, the case, for when both claws were reflexly exercised neither formed a crusher, signifying rather that bilateral differences in predominantly mechanoreceptive input to the paired claws somehow lateralized the claw ganglion [central nervous system (CNS)] into a crusher and cutter side. The side experiencing the greater activity becomes the crusher side while the contralateral side becomes the cutter and is also inhibited from ever becoming a crusher. This initial lateralization in the CNS is expressed, via as yet unknown pathways, at the periphery in claw morphology, muscle composition, and behavior. The critical period defines a time when the CNS is susceptible to being lateralized into a crusher and cutter side. Such lateralization is dependent upon experience of the environment in the form of mechanoreceptive input. In the absence of such experience, the CNS is not lateralized and paired cutter claws develop. Thus, while the critical period for crusher determination is genetically determined the actual trigger is influenced by experience. © 1992 John Wiley & Sons, Inc.     

Functional significance of an unusual chela dimorphism in a marine decapod: specialization as a weapon?

The squat lobster Munida rugosa has an unusual chela dimorphism exhibited mainly by large males. Some individuals have 'arched' chelae in which there is a gap between the dactylus and the pollex when closed, and others have a 'straight' morphology in which the dactylus and pollex oppose along most of their length. Geometric morphometric analysis indicated that, compared with males, the arched morphology does not develop fully in females, so further investigation was confined to males. In males, the distal part of the chela was similar in both the forms and seemed to be adapted to hold and shred prey items. Both morphologies had a major cylindrical tooth on the inner proximal part of the dactylus, but the arched morphology had a higher and wider propodus, a greater major tooth-pollex distance and a greater force generation than the straight morphology. The findings suggest that the arched chela morphology in M. rugosa is a sexually selected trait adapted to inflict puncture wounds on opponents during agonistic interactions. The arched morphology, therefore, appears to have evolved in males by means of sexual selection because it enhanced the function of the chela as a weapon, while retaining functionality for feeding.     

The defensive role of scutes in juvenile fluted giant clams (Tridacna squamosa)

This study tests the hypothesis that the scaly projections (scutes) on the shells of juvenile giant fluted clams, Tridacna squamosa, are an adaptation against crushing predators such as crabs. The forces required to crush scutes and clams were measured with a universal testing machine whereas crab chela strength was measured with a digital force gauge connected to a set of lever arms. Results for shell properties and chela strength are used to create two, non-mutually exclusive, predator–defense models. In Model 1, scutes increase the overall shell size, consequently reducing the number of crab predators with chelae that are large enough to seize and crush the prey. In Model 2, the chela has to open more to grasp a prey with these projecting structures which leads to a loss of claw-closing force such that crabs fail to crush the scutes, and consequently the clam. Clam scutes may also deter crab predators by increasing the risk of claw damage and/or handling time.     

Crab-resistant features of shells of burrowing bivalves: Decreasing vulnerability by increasing handling time

The vulnerability of burrowing bivalves to shell-breaking predation by crabs was found to be influenced strongly by shell features: size, shell thickness, degree of inflation, and the presence or absence of a gape.The relationship between the critical size of a bivalve (maximum size of vulnerability) and crab size was determined for four different morphotypes of bivalves. For the three bivalves where a “size refuge” was present, critical size increased with crab size. Nevertheless, when offered a choice crabs preferred clams well below the critical size and ate them in the order predicted by the critical-size experiment.Examination of the mechanics of shell crushing revealed how these shell features decreased vulnerability. Larger crabs could efficiently handle larger clams because both chela strength and degree of chela gape increased with crab size. Strain gauges attached to crab chelae showed that thick-shelled clams resisted a greater total number of force pulses than did thin-shelled clams of the same body weight. This may be related to the ability of thick-shelled clams to withstand greater loads than thin-shelled clams when loaded only once. This suggests that the reason for the increased resistance to crabs is prolongation of the shell-breaking time. Even though a large thick-shelled, tightly-closing, clam could eventually be opened, it will probably be rejected in favour of prey with shorter handling times.     

The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

The mechanics of predation by the shore crab,Carcinus maenas (L.), on the edible mussel,Mytilus edulis L.

Summary Mechanical aspects of predation by the shore crab, Carcinus maenas, on the edible mussel, Mytilus edulis, were examined. The shore crabs from the population studied utilized five distinct, largely size-related, mussel-opening techniques. Crushing the mussel umbone appeared the most successful opening method for medium-sized prey. Small mussels were crushed outright and large mussels could be opened by a slow, uneconomical, boring technique. The strengths of mussels, from an exposed shore, were tested under compression in four separate planes to determine the loads a crab would need to apply to crush the shells outright and the mechanical properties of mussels. Little inter-plane variability in compressive strength was observed, although intra-plane variability appeared high. The compressive strengths of mussels from a sheltered shore were found to be significantly higher than those from the exposed shore in the plane tested. A strain gauge was embedded in a mussel shell enabling the pattern and magnitude of forces produced by crab chelae in opening a mussel to be studied. The crab's chelae did not appear overwhelmingly strong when compared directly to the compressive strength of the crab's preferred mussel sizes. It is, therefore, postulated that crabs usually seek out and exploit weak spots in the umbone of mussels by trial and error, eventually breaking through the shell by a cumulative process of extending minute fractures in the shell substructure.     

Prey-induced changes to a predator's behaviour and morphology: Implications for shell-claw covariance in the northwest Atlantic

Whereas many plasticity studies demonstrate the importance of inducible defences among prey, far fewer investigate the potential role of inducible offences among predators. Here we ask if natural differences in a snail's shell hardness can induce developmental changes to a predatory crab's claw size. To do this, we fed Littorina obtusata snails from either thick- or thin-shelled populations to captive European green crabs Carcinus maenas. The crabs' shell-breaking behaviour dominated among those fed thin-shelled snails, whereas crabs fed thick-shelled snails mostly winkled flesh through the shell opening without damaging the shell itself (a.k.a. aperture-probing behaviour). Significantly, the size of crab crusher claws grew in proportion to the frequency of shell-crushing behaviour and, for a same shell-crushing frequency, crabs fed thick-shelled snails grew larger crusher claws than those fed thin-shelled snails after two experimental moults. Diet and behaviour had no effect on the growth of the smaller cutter claws of same individuals, providing good evidence that allometric changes to crusher claws were indeed a result of differential use while feeding. Findings indicate that both predation habits and claw sizes are affected by green crabs' diet, supporting the hypothesis that prey-induced phenotypic plasticity contributes to earlier accounts of shell-claw covariance between this predator and its Littorina prey in the wild.     

Can morphological and behavioral traits predict the foraging and feeding dynamics of social arachnids?

Complex social insect species exhibit task specialization mediated by morphological and behavioral traits. However, evidence of such traits is scarce for other social arthropods. We investigated whether the social pseudoscorpion Paratemnoides nidificator exhibits morphologically and behaviorally specialized individuals in prey capture. We measured body and chela sizes of adult pseudoscorpions and analyzed predation processes. Larger individuals spent more time moving through the colony and foraging than smaller pseudoscorpions. Individuals that captured prey had increased body and absolute chelae sizes. Although larger individuals had relatively small chelae size, they showed a higher probability of prey capture. Larger individuals manipulated prey often, although they fed less than smaller pseudoscorpions. Individuals that initiated captures fed more frequently and for more time than the others. Natural selection might be favoring individuals specialized in foraging and colony protection, allowing smaller and less efficient adults to avoid contact with dangerous prey. To our knowledge, there is incipient information regarding specialized individuals in arachnids, and our results might indicate the emergence of a morphologically specialized group in this species.     

Ultrasound-based subject-specific parameters improve fascicle behaviour estimation in Hill-type muscle model

The estimation of muscle fascicle behaviour is decisive in a Hill-type model as they are related to muscle force by the force–length–velocity relationship and the tendon force–strain relationship. This study was aimed at investigating the influence of subject-specific tendon force–strain relationship and initial fascicle geometry (IFG) on the estimation of muscle forces and fascicle behaviour during isometric contractions. Ultrasonography was used to estimate the in vivo muscle fascicle behaviour and compare the muscle fascicle length and pennation angle estimated from the Hill-type model. The calibration–prediction process of the electromyography-driven model was performed using generic or subject-specific tendon definition with or without IFG as constraint. The combination of subject-specific tendon definition and IFG led to muscle fascicle behaviour closer to ultrasound data and significant lower forces of the ankle dorsiflexor and plantarflexor muscles compared to the other conditions. Thus, subject-specific ultrasound measurements improve the accuracy of Hill-type models on muscle fascicle behaviour.