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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.

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., (© 2015. Published by The Company of Biologists Ltd.)

Hatching of embryos in the estuarine crab Sesarma haematocheir is highly synchronized with nocturnal high tide and completes within 1 hr among all embryos incubated by the female. This highly synchronized hatching is induced by a "Hatching-Program Inducing Factor (HPIF)" released from the female. To further define the cues involved in synchronized hatching, experiments were designed to characterize this factor and to determine possible sites of release and temporal release patterns using strategies involving isolation of egg masses, perfusion, and ablation experiments on fully developed embryos that had not yet entered the hatching program. Embryo transplantations indicate that not only HPIF may be released from the branchial chamber, but that it is extraordinarily unstable, and loses activity within 15 min, which frustrates further attempts at characterization. Nevertheless, with regard to temporal release patterns, it was established that HPIF activity was detected during short periods over three consecutive nights prior to release of larvae. This activity did not explain the gated response of embryo release in this crab, which might correspond with circatidal larval release events in the field., ((c) 2006 Wiley-Liss, Inc.)

The egg attachment system of an estuarine crab Sesarma haematocheir is formed on the maternal ovigerous hairs just after egg laying, and slips off these hairs just after hatching. The stripping is caused by an active factor that we call OHSS (ovigerous-hair stripping substance), which is released by the embryo upon hatching. OHSS was purified, and its active form had a molecular mass of 25 kDa. The cDNA of OHSS cloned from an embryonic cDNA library was 1759 bp long, encoding 492 amino acids in a single open reading frame (ORF). The C-terminal part of the predicted protein was composed of a trypsin-like serine protease domain, with homology to counterparts in other animals of 33-38%. The predicted protein (54.7 kDa) secreted as a zymogen may be cleaved post-translationally, separating the C-terminal from the N-terminal region. The OHSS gene was expressed in the embryo at least 2 weeks before hatching. Expression was also detected in the zoea larva 1 day after hatching and in the brain of the female. However, it was not detected in the muscle, hepatopancreas or ovigerous seta of the female. Ultrastructural analysis indicated that the material investing maternal ovigerous hair, i.e. the outermost layer (E1) of the egg case, is attached at the special sites (attachment sites) arranged at intervals of 130-160 nm on the hair. It is suggested that OHSS acts specifically at these sites, lysing the bond with the coat, thus disposing of the embryo attachment system. This enables the female to prepare the next clutch of embryos without ecdysis.

A pair of rosette glands (one of the tegumental glands in crustaceans) is present at the root of the dorsal spine of the thorax in mature embryos of the estuarine crab Sesarma haematocheir. Each rosette gland is spherical, 45-50 microm in diameter. This gland consists of three types of cells: 18-20 secretory cells, one central cell, and one canal cell. The secretory cells are further classified into two types on the basis of the morphology of secretory granules. There are 17-19 a cells, and only one b cell per rosette gland. An a cell contains spherical secretory granules of 2-3 microm in diameter. The granules are filled with highly electron-dense materials near the nucleus but have lower electron-density near the central cell. The secretory granules contained in the b cell have an irregular shape and are 1-1.5 microm in diameter. The density of the materials in the granules is uniform throughout the cytoplasm. The secretory granules contained in both the a and b cells are produced by the rough endoplasmic reticulum. Materials in the granules are exocytotically discharged into the secretory apparatus inside the secretory cell, sent to the extracellular channels in the central cell, and secreted through the canal cell. The rosette gland can be distinguished from the epidermal cells 2 weeks after egg-laying and the gland matures just before hatching. Materials produced by this gland are secreted after hatching and secretion continues through five stages of zoeal larvae. These rosette glands were never found in the megalopal larva. Rosette glands are found in the embryos of Sesarma spp. and Uca spp. In other crabs, tegumental glands are also found at the same position as in the embryo of S. haematocheir, but the fine structure of their glands is largely different from that of the rosette gland. On the basis of the morphology of secretory cells (a-g cell types), the tegumental glands of a variety of crab embryos can be classified into four types, including rosette glands (type I-IV). The function of these tegumental glands is not yet known, but different types of the gland seem to reflect the phylogeny of the crabs rather than differences of habitat., (Copyright 2003 Wiley-Liss, Inc.)

Embryos attached to the female crab Sesarma haematocheir hatch synchronously within 1 h. Hatching is also synchronized near the time of the expected nocturnal high tide. These events are governed by a single circatidal clock (or pacemaker) in the female crab. The present study examined the role of the optic peduncle of the eyestalk on hatching and hatching synchrony. Surgery was performed either from the tip of the eyestalk [to remove the region of the optic peduncle from the compound eye-retina complex to the medulla interna (MI)] or from a small triangle 'window' opened on the eyestalk exoskeleton [to create lesions on the medulla terminalis (MT) of the optic peduncle]. Neither hatching nor hatching synchrony was affected by removal of the region of the optic peduncle from the compound eye-retina complex to the MI: the circatidal rhythm also remained. Removal of the MI probably caused damage to the sinus gland and the bundle of axons running from the sinus gland to the X organ. Nevertheless, maintenance of highly synchronized hatching indicates that the X organ-sinus gland system is not related to hatching. Hatching and hatching synchrony were not affected by dorsal-half cuts of the MT: the timing of hatching was not affected either. By contrast, transverse and ventral-half cuts of the MT caused severe damage to most females; hatching of many females was suppressed, while hatching of some females was either periodic, at intervals of approximately 24 h, or arrhythmic for a few days. The bundle of neuronal axons is tangled in the MT, and the axons inducing hatching pass through the ventral half of the MT. Complete incision of these axon bundles may have suppressed hatching. Incomplete incision of the axon bundle or partial damage to the neurons may have caused periodic or arrhythmic patterns of hatching. There are two possible roles for MT in hatching. One possibility is that neurons in the MT only induce hatching under the control of the circatidal pacemaker located in a site somewhere other than the optic peduncle. Another possibility is that the circatidal pacemaker is actually present in the MT. The second possibility seems more plausible. Each embryo has a special 48-49.5 h developmental program for hatching. This program could be initiated by the circatidal pacemaker in the female, and hatching synchrony may also be enhanced by the same pacemaker.