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BILATERAL GYNANDROMORPH of the FRESH-WATER CRAB POTAMON FLUVIA TILE HERBST (DECAPODA: BRACHYURA) Fiorenza Micheli

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BILATERAL GYNANDROMORPH of the FRESH-WATER CRAB POTAMON FLUVIA TILE HERBST (DECAPODA: BRACHYURA) Fiorenza Micheli !"#$%&'$#()*+$+,'-.-'/0(-1(%0&(2'&3045$%&'(6'$7(8-%$.-+(1#9:"$%"#&(;&'73%(<=&>$/-,$? !'$>0*9'$@ A9%0-'<3@?(2"-'&+B$(C">0&#" D&:"&E&,(E-'F<3@? G-9'>&?(H-9'+$#(-1(6'93%$>&$+(!"-#-I*J(K-#L(MMJ(N-L(O(<N-:LJ(MPPM@J(//L(QRM4QRS 897#"30&,(7*?(The Crustacean Society G%$7#&(TDU?(http://www.jstor.org/stable/1548526 . A>>&33&,?(VQWVXWYVMY(YV?MY Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The Crustacean Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Crustacean Biology. http://www.jstor.org JOURNAL OF CRUSTACEAN BIOLOGY, 11(4): 561-568, 1991 BILATERAL GYNANDROMORPH OF THE FRESH-WATER CRAB POTAMON FLUVIA TILE HERBST (DECAPODA: BRACHYURA) Fiorenza Micheli ABSTRACT The morphologyand gonad history of a bilaterallygynandromorphic fresh-water crab (Po- tamonfluviatileHerbst) is described.The abdomen and gonopods exhibit male characteristics on the left side of the body, female ones on the right,with the exception of a partialmasculin- ization of the first right pleopod. The right side has a normal female gonopore, the left one a penis. Internally,testes and vas deferensare presenton both sides, thoughthe rightvas deferens is incomplete and does not communicatewith the outside. On the right, a seminal receptacle full of sperm indicates that the animal has mated as a female. An isolated portion of gonad, also showingtesticular activity, is annexedto the seminal receptacle.Mature sperms are visible throughoutthe gonadand vas deferens.No ovariantissue was detectedin the entirereproductive apparatusof this specimen. The haploid chromosome number does not differ significantly betweenthe male and female sides and averages68 ? 1.3. The role of the androgenichormone on sex differentiationof Brachyurais discussed. Though bilateral gynandromorphism its role is uncertain in the Reptantia. In among normally dioecious decapod crus- decapods, the AG is thought not to initiate taceans has already been reported (Chace sex differentiation,though it probably ini- and Moore, 1959; Ito, 1965; Farmer, 1972; tiates puberty(Charniaux-Cotton, 1975). In Otto, 1979; Cargo, 1980; Johnson and Otto, several cases, implantation of the AG into 1981; Manningand Holthuis, 1981; Taylor, prepubertaland pubertal decapod females 1986), few works describe the gonad his- did not masculinize the ovaries and seldom tology (Farmer, 1972; Cargo, 1980; John- masculinized the external sexual character- son and Otto, 1981). It is evident from the istics (Charniaux-Cottonand Payen, 1988). literatureon this topic that various degrees In contrast with these results, the sex re- of sexual differentiationhave been encoun- versal of nonhermaphroditicfemale prawns tered on the two sides of such specimens, (both matureand immature)implanted with rangingfrom definitebilaterality in both the the AG seems to indicate that this gland has primaryand secondarysexual characters (the a similar function in Amphipoda, Isopoda, lobster Homarus americanus, Chace and and Decapoda (Nagamine et al., 1980a, b). Moore, 1959; the fresh-waterprawn Neph- A population of Potamon fluviatile in- rops norvegicus,Farmer, 1972), to either in- habiting a hill stream near Florence, Italy complete development (the blue crab Cal- (see referencesbelow for a descriptionof the linectes sapidus, Cargo, 1980) or partial study site), has been the object of extensive masculinizationof the gonad on the female ethological and ecological studies (Gherar- side (Callinectessapidus, Johnson and Otto, di, 1987), including reproductive biology 1981). and growth patterns (Micheli et al., 1990). Such abnormalsexual differentiationsare Of the approximately2,000 crabs sampled of interest for what they may contribute to or observed in the course of 10 years' re- understanding the mechanisms of sexual search,only one sexuallyabnormal individ- determination, well known in some crus- ual was encountered.This paperreports the tacean groups such as Isopoda and Am- morphologyand histology of this specimen. phipoda (Charniaux-Cotton and Payen, 1988), but still debated within Decapoda. MATERIALSAND METHODS In particular,while the role of the andro- The gynandromorphwas collected in August 1988, genic gland (AG) in determiningthe differ- taken to the laboratory,and killed with chloroform. entiation of and sexual All its internal organs were normal in size and ap- primary secondary pearanceexcept for the gonads, which were removed, charactershas been proven in the former fixed in Carnoy'sfluid, and preservedin 75%ethanol. two groups and in the Decapoda Natantia, No evidence of a pathologiccondition or of parasitic 561 562 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991 Fig. 1 Ventral view of a bilateral gynandromorph of Potamonfluviatile, showing the anomalous abdomen and gonopods. Male characteristics are evident on the left side of the body, female ones on the right side. infestation was detected. The whole gonad was divided (Fig. 1). The abdomen was asymmetrical, into subsamples and, along with the seminal receptacle, broad and rounded on the right (female) embedded in polystyrene resin, serially sectioned (7 narrow and on the left. The Am), and stained with hemalum-eosin. side, straight Meiotic chromosomes were prepared following a two typical male pleopods were present on technique suggested by I. Lazzaretto and A. Libertini the left side. On the right, the second, third, (personal communication). Immediately after dissec- and fourth pleopods were biramous, with tion, tissue samples taken from both the left and right setae, as in normal females, while the first sides of the gonad were kept in bidistilled water for 10 min, then fixed in 3:1 ethanol/acetic acid and preserved was also biramous, but shorter and thicker at 4?C. Tissue subsamples were placed in 60% acetic and with no setae, thus slightly masculin- acid for 3-5 min, then broken up by pipetting and ized. One female gonopore was present on ejecting the liquid and tissue several times. One drop the ventral right side of the sixth thoracic of this cellular suspension was placed on a slide and one on the coxa of the left stained in Giemsa diluted with a phosphate buffer 0.15 segment, penis M (1:20). fifth pereiopod. The gonads appeared normal on the male RESULTS side; testis and vas deferens were complete and well developed as in a mature male. Anatomy The dorsal portion of the gonad on the fe- The crab had a carapace length (CL) of male side was similar to a testis, being white 36.8 mm. It was, therefore, an adult, since, and slender instead of thick and orange as in this species, the puberty molt occurs in normal females of the same size. This around 35 mm CL (Micheli et al., 1990). testis was slightly smaller than the left one. The histological examination confirmed that A transverse commissure, leading to the go- this specimen had mature gonads and there- nad on the other side, was present. fore had undergone the puberty molt (see A vas deferens had developed on the right below). The crab had a hard exoskeleton, side as well, but did not lead to the right no sign of scars or mutilation, and was right- genital opening. A small, unbranched por- handed. tion of gonad, white like the rest of the re- Externally, abnormality was evident only productive apparatus, was present on the in the shape of the abdomen and pleopods right ventral side. This part of the gonad MICHELI: GYNANDROMORPHISM IN FRESH-WATER CRAB 563 ,I?, ?r?~~~.q.; 1 ?urTl 'A ' s '~ ,, * w _dM ?/ - <-? L- *l Tfc. '' f^ i , , t; . , ??-< 7-.*.? ,i9 *~~~S * If~ * ' L^.j# t , ? " ~, I'w..- ' ., i? t 'IH, v- ' .i &- Fig. 2. Histological sections of the gonads of a bilateral gynandromorph of Potamonfluviatile. Hemalum-eosin. C = connective tissue. S = spermatocytes. Sp = sperms. a, Testicular lobules on the male side. b, Testicular lobules in the dorsal portion of the gonad on the female side. c, Testicular lobules in the portion of the gonad on the female side annexed to the seminal receptacle. d, Magnification of testicular lobules on the female side (as in b). Connective tissue separates lobules containing cells in the same maturational stage. 564 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991 was connected to an apparently normal The seminal receptacle resembled normal seminal receptacle and to the female gon- ones, previously sectioned (Micheli et al., opore. 1990). The wall consisted of a monolayered, No connection was observed between the tall columnar epithelium resting on a con- ventral and dorsal gonads on the right side. nective tissue sheath. Sperm masses were visible in the lumen (Fig. 3b), indicating Histology that the animal had mated as a female. All the gonad portions were masculin- Several meiotic divisions were observed ized. Testicular lobules constituted the dor- on slides prepared for chromosome exam- sal portions of both the right and left gonads ination. Pachytene stages prevailed. Unfor- (Fig. 2a, b, d), and also the ventral portion tunately only a few prometaphase plates, of the right gonad, annexed to the seminal characterized by short, condensed chro- receptacle (Fig. 2c). In the isolated ventral mosomes (Fig. 4), proved suitable for chro- portion of the right gonad, abundant, poorly mosome counts. In all other stages, chro- structured connective tissue filled the inter- mosomes overlapped extensively and could lobular spaces (Fig. 2c). not be individually distinguished. In the few All stages of spermatogenesis, mature plates selected, chromosomes were so vari- sperms included, were detected in the go- able in length and shape that individual nad. Cells in the same developmental stage chromosomes were not recognizable and a seemed to aggregate within the same lobules karyotype could not be made.
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