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Chiromantes (Brachyura: Sesarmidae)

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Chiromantes (Brachyura: Sesarmidae) Bio logy of Anomura II (A. Asakura，e d.) ，Crustacean Research，Specia lNu mber6: 109-119，2006 Expression and structure of stress chaperon hsp90 in terrestrial decapods，Coenobita (Anomura: Coenobitidae) and Chiromantes (Brachyura: Sesarmidae) Oleg Gusev，Tracy A. Ziegler andMasayuki Saigusa A bst ract .- Thisstudy describes aspects restrial adaptation. Both Coenobita spp.and of the biologyof heat shock proteins 90kDa Chiromantes haematocheir b巴long to the T4 (hsp90) in terrestrial decapods，Coenobita pur- group，since they are fo und in aterrestrial envi ・ pureus，C. bre vimanus (Anomura: Coenobitidae) ronment as adult s，but still depend on the ma- and Chiromantesha ematocheir (Brachyura: rine environment for planktonic larval develop- Sesarmidae). These species possess highlydevel- m en t. oped terrestrial adaptations on both morphologi- Recent studies of the concomitant effects of cal and phys iological levels. We have analyzed therrnal acclimation onboth heat and cold tol - the pattern of express ion for th ege ne coding erance have been made in m anyspecies: e.g . hsp90:mol ec ular chaperon in both embryos and ， crabs (Sti llm an 2003) planarians (Ts ulωda the muscle tissues of aduIt sof these species in ， ， re sponse to heat stress. While the minimum tem- &Ogoshi ，19 78) ，earthworms (Hanumante， perature for initiation of the synthesis of excess 1977) ，flies (Ohtsue tal. ，1999) ，lobsters (Spees of hsp90 mRNAwas different between embryos et al. ，2003) ，fish (Hemandez & Buckle，2002) ， and aduIt sfrom the different climatic region s， crayfish (Layne et al. ，1985) ，and copepods the range of re sistance to high temperature shock (Bradley，1 978; Voznesens)くy etal .，2004) . was nearl ythe sa me (6 ーやり. Furthermore，the Taken to gether，the res ult ss uggest that ，even str ucture of the stre ss -factor binding domain in if they are close systematica l1 y，or ganisms hsp90 codin ggene showed convergent pattern sb y exposed to the same climatic conditions will possessing aconservative amino acid se quence in display different bodytemperatures ，effective decapods experiencing the sa me annual tempera- metabolic temperatur es ，and patt emingof heat ture regimes in the subtropical，te mperate and stress re sponse and ac climation (He1mut et a l.， boreal climatic regions. 2002). Compared to other arthropods，terrestria1 decapod crustaceans have arelati ve ly10 w re- INTRODUCTION sis tance to hi ghtemper at ure stress and water Two separateIi nesof decapod crustaceans 10ss (Schmitz & Harrison ，2004). These two have successfully in vaded the terrestrial envi- pr oblemswere overcome by the deve10pment ronment (for review ，see Schmitz & Harrison， of physio10 gica1 and behavioral adaptations ， 2004) the hermit crab family Coenobitidae including dai 1ymi gration s(I mafuku，2002; (Anomura) (for review ，see Burggren & Morris，2002) ，comp li cated breedingand 1ar- McMahon，198 8; Greenaway，2003) and vae release behaviors (Saigusa & Terajima， the Grapsoidea (Brachyura) (Adamczewska 2000;Im afuku，2002; Brodie，2002) ，changes & Morris，2000; Anger & Shubart，2005). in respiration ，in circu1atory systems，and in Decapods th at spend more or less of their time the dynamicsof molting and food di gest ion in the terrestrial environment have been sum- (McMahon & Burggren，1988 ;Morris ，2002) . marized into five grades，T 卜T 5 (Burggen & An important question is whether evo1u- McMahon，199 8) ，according to degree of ter- tionary adaptation at phenotypic and genotypic 110 O. Gusev efal levels affects acclimation and general com- the highest temperature and water level varia- pensatory abilities of terrestrial crustaceans to tion ，were shown to possess three broad catego- environmental temperature changes.Nearly ries of variation of response. First ，total cellular all members of Coenobitidae use behavioral levels of hsp's varied with season (Hofmann & means to avoid overheating，but the tempera- Somero，1995; Buckley eta l.， 2001) and with ture threshold for such behavior was shown laboratory acclimation (Tomanek & Somero， to be higher in diumally active species，rather 1999; Buckley etal .，2001). Second，although then in nocturnal and more terrestrial species the stress response displays an element of plas- (Burggren & McMahon，1988; Greenaway， ticity，some components appear to be fixed， 2003). Meostatic temperature of the hemo- and are acharacteristic of aparticular species. Iymph in terrestrial hermit crabs and brachy- Different sets of congeners of rocky intertidal uran crabs is generally higher then in their lit- marine invertebrates have different stress re- toral relatives ，while the oxygenation level does sponses despite being acclimated to the same not show aremarkable difference (Burggren temperature (e.g .，Hofmann & Somero，1995; & Mahon，1988). Furthermore，the preferred Tomanek & Somero，2002). Third，the stress average body temperature and the temperature response varies in natural populations and close of hemolymph，as well as total hemolymph species across environmental gradients and oxygen capacity shows atendency to be higher with thermal habitat (Helmuth & Hofmann， in terrestrial rather than in marine species 2001). Moreover，there is alink between the (Burggren & McMahon，1988; Schmitz & structure of certain domains in major groups Harrison，2004). of constitutively expressed and stress-induced The question of genetypic basements for the chaperons and temperature-adaptive molecular physiological plasticity during heat acclimation response in the species with different latitudi- in crustaceans was partially answered using nal biogeographic patteming (Horowitz，2001; temperature-stress speci fi. cp 附 eins (molecular Somero，2005 ;Mahroof ，et al .，2005). chaperons) as markers of the compensatory In this study，we investigated the effect of abilities of the organism to overcome dam- temperature on the viability and expression age caused by excess environmental stress on of the molecular chaperon hsp90，showing acellular leve l. Heat shock proteins (hsp) are mixed cytosolic and stress-response nature of the most studied group of molecular chaperons the adults and embryos of two species of land involved in the refolding of proteins damaged hermit crabs Coenobita and abrachyuran crab by external stresses (Soti & Scermely，2003; Chiromantes haematocheir. To analyze the pos- Robert，2003). This group includes four m 勾or sible influence of primary and secondary struc- types of proteins (hsp27，hsp70 ，hsp90 and ture of hsp90 on the induction of increasing hsp 100) ，according to their molecular weight levels of this protein in response to heat shock， and functions (Feder & Hoffman，1999). Hsps we cloned anucleotide sequence coπesponding are particularly significant as they represent a to the particular region in the second functional mechanism by which an organism can buffer ATP-dependent domain of hsp90.This is one the impact of environmental temperature on the of the potential markers of environmental- protein pool without having to employ special- conditions based on changes in the functional ist protein isoforms to withstand high tempera- structure of hsp90 (Schnaider etal .，1999; Zhao tures (Somero，1995). The investment in hsp' et al. ，2005). This domain is regarded as the s，as the likelihood ofhigher temperatures from binding site for the heat shock factor (hs f) ，a global climate change looms，may be an e汀ec- protein controlling chaperon activity of hsp90 tive strategy for intertidal and terrestrial crusta- and intensity of its synthesis under stress con- ceans ditions .The activity of hsf closely depends on Crustaceans and some other inhabitants of the level of damage caused by high temperature the intertidal zone，which is characterized by and other stresses (Westwood & Wu，1993; Stress protein hsp90 in terrestrial decapods lll Rabindran et a l.， 1994).We also analyzed the Samples of boreal species，H yas araneus primary genetic structure of the heat factor (8rachyura: Maj idae) ，Sclerocrangon boreas binding site in hsp90 coding gene in anumber (Caridea:Crangonidae) and Pagurus pubescens of decapod crustaceans from different tempera- (Paguroidea: Paguridae)，were collected by ture regimes within various climatic regions. deep creeping (depth - 80-100 meters) in the Kandalaksha Gulf，White Sea，Russia (32-340E ， 65-660N).The water temperature at the collect- MATERIALS ANDル1 E THODS in gpoint depth was 4oC and thi svalue remains Field sampling constant throughout the year at these depths. No The main target species Coenobita purpu- specific acclimatization procedures were further reus，C. brevimanus and Chiromantes haema- applied to the adults and the embryos. tochei r， as well as the other decapod species for comparative purposes，were collected as fol- Heat stress treatment lows 什omthree regions in Japan with different For the excess temperature treatment，eight annual temperature fluctuation in both water rep1icate experiments were conducted. The em・ and air . bryo clusters (at the stage of development close Subtropical species were collected at to hatching; weight - 1g; 1200- 1300 embryos) lriomoteサima Island，Ryukyu (124 oE ，24 0N)， were dissected from the abdomen of females of with daily fluctuation of water temperature of Chiromantes and Coenobita and immediately about 24-260C on the surface and in shallow placed in a1.5 ml tube，which was preheated to water and air temperature of about 28-30 oC the experimenta1 temperature and filled with ar- during the season of collection (April-June， tificial sea water (10%0). Temperature treatments 2004-2005). Ovigerous females of land hermit
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