Shell Layers and Structures in the Brackishwater Bivalve, Corbicula Japonica
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ΐῒ῏ῑῐ ῍The Quaternary Research῎ ./ ῍ / ῎ p. -+1ῌ--+ Oct. ,**0 Shell Layers and Structures in the BrackishWater Bivalve, Corbicula japonica Keiko Yamaguchi* +,a, Koji Seto* ,, Katsumi Takayasu* - and Morihiro Aizaki* + Bivalve shells contain a great deal of information about the animal’s growth history and local environment. Opaque and translucent layers are observed in the outer shell of Corbicula japonica. Detailed imaging with scanning electron microscopy and laser microscopy show that microstructural features are di#erent between the two layers. The di#erence is mainly determined by the content of organic materials ; the translu- cent layer contains less organic matrix than the opaque layer. A marking experiment revealed that the translucent layers were formed in the period from early summer to winter, varying between individuals. If there is no formation of an opaque layer and shell growth rates are low in autumn, the translucent layer is accompanied by an annual ring on the shell surface. The period of formation of the layers is synchronized with the annual cycles of stable isotope values, especially with oxygen (d+2O). By combining the observation of the shell structure with stable isotope analysis, the shell of C. japonica can be used to provide information for reconstructing ecology and estuarine environments in the past. Keywords : brackish water, bivalve, Corbicula japonica, microstructure, shell growth, translucent layer, stable isotopes their response to environmental variation (e.g. I. Introduction Carter, +32* ; Dillon and Clark, +32*). Shell Bivalve shells can contain a detailed record microgrowth patterns are useful for recon- of the animal’s growth history and environ- structing tidal cycles or sea-level in the time and ment, similar to other accretionary growth skel- place the animal lived (e.g. Lutz and Rhoads, etons, such as corals or tree rings. The infor- +32* ; Ohno, +323). Compared to many other car- mation stored within a bivalve’s hard parts is bonate skeletons (e.g. foraminifera, ostracods, very useful for the reconstruction of environ- nannofossils), the bivalve shell is very well mental changes. It is well known that bivalve suited for the reconstruction of detailed time shell chemistry holds a record of environmental series of environmental change covering the change encountered by the animal. The car- life of the animal. bonate shell can provide a record of oxygen Shell also contains a great deal of informa- isotope variations (d+2O) that is often used as tion on the age and growth properties of the paleo-thermometer (e.g. Craig, +30/ ; Rye and animal. Age determination based on annual Sommer, +32*). Growth-line analysis is also rings on the shell surface has been carried out recognized as a useful method to investigate in many marine bivalves (e.g. Orton, +3,0 ; Naka- the life history and growth rates of clams and oka, +33,). An annual ring is, in many cases, a Received April 1, ,**0. Accepted June +3, ,**0. This paper was presented on the Symposiumofthe,**/th Annual MeetingoftheJapanAssocoationfor Quaternary Reaeach. * + Faculty of Life and Environmental Science, Shimane University. +*0* Nishikawatsu, Matsue, 03*ῌ2/*.,Japan. * , Research Center for Coastal Lagoon Environments, Shimane University. +*0* Nishikawatsu, Matsue, 03*ῌ2/*.,Japan. * - Shimane University. +*0* Nishikawatsu, Matsue, 03*ῌ2/*.,Japan. *a Corresponding author:[email protected]. ac. jp 318 Yamaguchi, K., Seto, K., Takayasu, K. and Aizaki, M. Oct. ,**0 slow-growth zone formed when the shell growth served that the annual ring is associated with stopped in winter (winter break). The spawn- the translucent layer within the mineralized ing break in shell growth is also recognized shell of C. japonica, collected from Lake Aba- in many marine bivalves (e. g. Rhoads and shiri. He recognized that the shell of C. japonica Pannella, +31* ; Sato, +33/). has two characteristic layers, translucent and Though marine bivalves have often been the opaque, and he suggested that the translucent objects of such studies, applying these tech- layer is formed when the shell growth rate is niques to brackish water bivalves has not been low. The translucent layer has been recognized common. As environmental variation associ- in other bivalves, e.g. Anadara (Kobayashi, +310), ated with climate change may be amplified at Gomphina (Kobayashi, +313), Mytilus (Hosomi, the boundary between land and sea, studies of +323), and been regarded as an annual marker. environmental records stored in the shell geo- Kobayashi (+310) reported that shell structure chemistry and microstructure of brackish bi- of the outer shell layer of Anadara broughtonii valves become more valuable. changes between crossed lamellar and compos- Corbicula japonica is an endemic bivalve as- ite prismatic structures alternately, and that sociated with the low salinity brackish environ- the crossed lamellar structure is rather trans- ment. It inhabits brackish lakes and estuaries lucent and formed accompanied with a slow- from Hokkaido to Kyushu in Japan, and is a growth zone. Hosomi (+323) observed white very important species for inland fisheries in rings corresponding to the translucent layer in Japan (Nakamura, ,***). Paleo-environmental shells of Mytilus galloprovincialis, and used them information can be provided from the shells of for age determination. But he did not describe C. japonica, not only from fossil deposits but the microstructure of the increment nor the also from shell middens associated with an- time of formation of the rings, and whether it is cient human habitation. related to environmental factors or not. Prezant Kobayashi and Takayasu (+33/) studied shell et al. (+322) investigated color change in fresh- microstructure of Japanese brackish-water cor- water Corbiculid shell layers, and argued that biculids in detail and observed that the outer color change occurs when an animal is exposed shell layer of C. japonica is composed of the com- to a significant stress. This change is between plex crossed lamellar structure (finely-crossed purple and white layers in the shell. Kobayashi type). Takayasu et al. (+330) carried out a mark- and Takayasu (+33/) also mentioned that the ing experiment to examine the process of mi- calcareous shell layers of Corbiculidae species crogrowth increments of C. japonica in Lake are composed of wider dark and narrower light Shinji, and found that the microgrowth pattern band arranged alternately. These probably cor- showed neither cyclicity nor periodicity. They respond to the translucent and opaque layers, suggested that the formation of microgrowth but there has been no detailed observation or increments was most likely influenced by com- description of these shell layers in the litera- plex changes in the brackish water environ- ture. ment as well as by physiological rhythms and Before shell layers can be used as the basis of individual variation in these physiological fac- studying variation in the brackish water envi- tors. ronment, more work needs to be done on shell On the contrary, several studies on shell structure and growth banding, particularly fac- growth and age determination of C. japonica tors controlling opaque and translucent layers. were carried out based on annual rings in shell The purpose of this study is to describe the (Utoh, +32+ ; Takada et al., ,**+ ; Oshima et al., structural di#erence between the opaque and ,**.). Although some specimens have clearly translucent layers and to document the forma- recognizable annual rings, it is usually di$cult tion of the two layers. Finally we will examine to identify annual rings on the shell surface, the possibility of using these layers for the age owing to the thick, dark colored periostracum determination, growth analysis, and paleo-envi- and the existence of irregular concentric rings ronmental reconstruction. covering the surface of the shell. Utoh (+32+)ob- ,**0 ῒ +* ῑ Shell layers in Corbicula japonica 319 are secondary emission images (SEI) taken at II. Materials and methods accelerating voltages of /4* or +* kV. A laser +. Materials microscope (KEYENCE VKῌ2/**) was used to Living individuals of Corbicula japonica were produce a -D image of the etched shell section. collected between December +333 and June ,**/ The outer shell layer of one sectioned valve from Lakes Shinji, Jinzai, and Nakaumi, eastern was sub-sampled at fine spatial resolution (*4. Shimane Prefecture, southwest Japan (Fig. +). mm thick layers) along the shell surface to ex- Specimens used in this study were about , cm amine the change of oxygen and carbon stable in length, typical of adult-size clams. They lived isotope values throughout growth and in rela- in shallow water along the coast of these brack- tion to shell layers. A gas-ratio mass-spectro- ish lakes. Because the tidal amplitude in these meter (Finnigan MAT delta S) was used to lakes is very small, clams remain covered by measure the oxygen and carbon stable isotope water at all times. ratios. Analyses were calibrated relative to The salinity of the water at the habitat of C. the PDB standard and the standard deviation japonica in Lake Shinji is about / psu. That of of repeated internal standards was less than Lake Jinzai is somewhat variable, between -῍ ῎*.*/ῐ. +2 psu. The salinity of the shallow water in -. Field experimental methodology Lake Nakaumi is higher, +* to ,* psu. The wa- A field experiment was conducted to analyze ter temperature ranged from , to -,῏ in all the the growth of the C. japonica and to test the lakes. timing of formation of the translucent layer. ,. Observation of shell structure Two artificial ponds were used for culturing A single valve from each specimen was sec- C. japonica beside Lake Jinzai. Brackish lake tioned from umbo to the ventral margin to ex- water was continuously supplied to the ponds. amine the shell structure. The sectioned valve Pond dimensions were +* m in length, / min was observed with a binocular microscope, and width and ,*῍0* cm in depth. the thickness of the layer formed each month Clams were distributed throughout one of was measured.