Ontogeny of Cheliped Laterality and Mechanisms of Reversal of Handedness in the Durophagous Gazami Crab, Portunus trituberculatus.

The paired claws in Gazami crabs, Portunus trituberculatus, are bilaterally asymmetrical, and asymmetry is remarkable on the distal two segments of the first pereiopod, that is, the dactylus and propodus. Shells are exclusively cracked by use of the right chela, representing handedness. In Gazami crabs, handedness is reversed after autotomy of the right chela. Our study focused on the ontogeny of handedness and the mechanism of handedness reversal. Morphologically, asymmetry was first detected in megalopa larvae where the right propodus was significantly larger than the left, as was the canine at the base of the right dactylus. Presumably, the rate of chelagenesis differed between the left and right chelae. With these morphological features, the right chela functioned as a crusher. The crusher exerted a closing force two to three times that of the cutter. With loss of the right crusher, the left chela was bigger than the regenerated right chela and was converted to the crusher. In contrast, the performance of the regenerated right chela deteriorated compared to that of the original right crusher, and exertion of full closing force was inhibited by the more active left chela. Furthermore, crabs with two crusher chelae did not clearly show handedness. A decrease in size and performance of the regenerated right chela can be explained by a default program hypothesis. In conclusion, a difference in the chelagenesis rate results in bilateral asymmetry of the two chelipeds, and then handedness is generated by neural regulation in the thoracic ganglion innervating these claws. Since handedness is reversed after autotomy, the thoracic ganglion would not be lateralized in Gazami crabs. A default program hypothesis is proposed to explain the ontogeny of bilateral chela asymmetry and handedness reversal.

Chela asymmetry in a durophagous crab: predominance of right-handedness and handedness reversal is linked to chela size and closing force.

The swimming crab Portunus trituberculatus is a durophagous brachyuran. Right-handed crabs are predominant, but left-handed crabs are also found in nature. Left-handedness may arise from loss of the right crusher. We examined whether heterochely (morphology) was correlated with differences in closing force (physical property) and handedness (behaviour). The closing force was stronger in larger chela with greater apodeme height and handedness resided in the chela with stronger closing force. With loss of the right chela (autotomy), handedness transitioned from the right to left chela, and all crabs were left-handed thereafter. Reversed handedness was accompanied with a reduction of size and closing force in the regenerated right chela, and growth of the original left chela. After handedness reversal, dentition on the left dactylus of the newly-converted crusher was close to that of the original right crusher, but did not attain the same shape, even after 10 moults. Left-handed crabs were significantly worse than right-handed crabs at crushing hard-shelled prey. Chela formation was symmetrical in the zoea, and heterochely and right-handedness started in the megalopa, regardless of maternal handedness. Since the left chela is capable of being the crusher, heterochely may be caused by differences in morphogenetic velocity between the right and left chelae, under a signal discriminating right from left. Right-handedness is an attribute of P. trituberculatus, that would be inheritable across generations. It is probable that right-handedness was used in the earliest durophagous crabs, and this trend has been succeeded to extant species.