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Juvenile hormone regulates extreme mandible growth in male stag beetles.$ ¥®ÂήɴÁªº¶Pº¸»X´xºÞ¤F¤½ÂΪº¤jÃEªº¦¨ªø!g Gotoh H, Cornette R, Koshikawa S, Okada Y, Lavine LC, Emlen DJ, Miura T.Y ¤@°ï¦W¦ry. ©©øÂν׾ -- ©øÂν׾¡@¡@ Cy Laboratory of Ecological Genetics, Graduate School of Environmental Science, Hokkaido University, Sapporo, Japan.[ ¥ÍºA¿ò¶Ç¬ã¨s«Ç,¤é¥»¥¾·E¥_®ü¹D¤j¾ÇÀô¹Ò¬ì¾Ç¬ã¨s©Ò!a@3Qk ©©øÂν׾ -- ©øÂν׾¡@¡@ ic Abstract¤jºõkWHKm ©©øÂν׾ -- ©øÂν׾¡@¡@ P%fyv The morphological diversity of insects is one of the most striking phenomena in biology.(P0[7W ©øÂΫ¬ºAºØÃþ°Ï¤À¬O¦b¥Íª«¾Ç¸Ì¨ä¤¤¤@¶µ³Ì¤Þ¤Hª`·Nªº¦ÛµM²{¶HJ ©©øÂν׾ -- ©øÂν׾¡@¡@ \ Evolutionary modifications to the relative sizes of body parts, including the evolution of traits with exaggerated proportions,are responsible for a vast range of body forms.2~a&QK ºt¤Æªº§ïÅܨìÓÅé¬ÛÃö¤Ø¤oªºÃö«Y,¥]§t¥H¸Ø±iªº¤jÃE¯S¼xªººt¤Æ¹Lµ{¬O¸ò¤j½d³òªº¨Å髬ºA¦³µÛ±K¤ÁªºÃö«Y.pq ©©øÂν׾ -- ©øÂν׾¡@¡@ ^ Remarkable examples of an insect trait with exaggerated proportions are the mandibular weapons of stag beetles.^ ¸û·|Åý¤Hª`·N¨ìªºÄ´¦p¦³µÛ¸Ø±i¤ñ¨Ò©øÂίS¼x.§â¤jÃE·íªZ¾¹ªºÁê§ÎÂÎLzh#F# ©©øÂν׾ -- ©øÂν׾¡@¡@ n6B Male stag beetles possess extremely enlarged mandibles which they use in combat with rival males over females. As with other sexually selected traits,
¤½Áê§ÎÂξ֦³·¥¤jªº¤jÃE,¥Î¦b¹ï§Ü¥ÀÂΨ®Çªº¨ä¥L¤½Áê§ÎÂιï¤â¤W,·í¥H¤jÃE·í¦¨©Ê§O¿ï¾Üªº¤@ºØ¯S¼x®É,hI stag beetle mandibles vary widely in size among males, and this variable growth results from differential larval nutrition.o ¦b©Ò¦³¤½Áê§ÎÂΨ½¤jÃE¦³«Ü¼e¼sªºÅܤÆ,³oÓ¦hÅܤƪº¥Íªø²£¥Í©ó¨Ó¦Û¤£¦Pªº¥®ÂήɴÁ¾i¤ÀT_Q3c' ©©øÂν׾ -- ©øÂν׾¡@¡@ ,Z; However, the mechanisms responsible for coupling nutrition with growth of stag beetle mandibles (or indeed any insect structure) remain largely unknown.nr[ µM¦Ó,¥æ°t¾i¤Àªº²Õ¦¨»PÁê§ÎÂΤjÃE¦¨ªø(©Î¥ô¦ó©øÂε²ºc)ªºÃö«Y¤´¦s¦b«Ü¤jªº¥¼ª¾¼Æ(y]MTi ©©øÂν׾ -- ©øÂν׾¡@¡@ w0 Here, we demonstrate that during the development of male stag beetles (Cyclommatus metallifer),
juvenile hormone (JH) titers are correlated with the extreme growth of an exaggerated weapon of sexual selection.M< ³o¸Ì,§ÚÌÃÒ¹ê...¦b¤½Áê§ÎÂΪºµo¨|´Á¶¡(¥H¬ü¥L¬°¨Ò)¥®ÂήɴÁªº¶Pº¸»X¶q¬O°Ï¤À©Ê§Oªº¸Ø±iªZ¾¹ªº·¥¤j¦a¦¨ªø¦³Ãö«Y)5P)*_
We then investigate the putative role of JH in the development of the nutritionally-dependent, phenotypically plastic mandibles,}}Fw] by increasing hemolymph titers of JH with application of the JH analog fenoxycarb during larval and prepupal developmental periods.
¦b¥®ÂΤεo¨|®É´Á,¹B¥Î¶Pº¸»Xªº¦¨¤ÀÀËÅ羯¤è¦¡¾aµÛ¼W¥[¥®ÂζPº¸»Xªº·»¾¯¶q,§ÚÌ©ó¬O¬ã¨s¥X¥®ÂζPº¸»X¦b¨Ì..¿à¾i¤Àªºµo¨|¦ÓÅãµÛ¦a§Î¦¨ªº¤jÃE©Ò§êºt¨¤¦â.nw ©©øÂν׾ -- ©øÂν׾¡@¡@ WyY, Increased JH signaling during the early prepupal period increased the proportional size of body parts, and this was especially pronounced in male mandibles,\ enhancing the exaggerated size of this trait.5 ¦bªì´Áªº¥®ÂήɴÁ,¤H¤u¼W¥[ªº¥®ÂζPº¸»X«H¸¹¦]¤l·|¼W¤j¤F³¡¤À¨Å骺¤ñ¨Ò(¥i¯à«ü¤jÃE),¨Ã¥B³o¦b¤½ÂΤjÃE¯S¼x¤W´£¤É¤F¸Ø±iªº¤Ø¤o¤W¦³¯S§O¦a©úÅã.
The direction of this response is consistent with the measured JH titers during this same period. Aa (¤Wz)¤ÏÀ³¥X¨Óªºµ²ªG¸ò¦b¦P¤@®É´Á¶q´ú¥Xªº¥®ÂζPº¸»X¾¯¶q¬O¦³¤@PªºÃö«Y.-/1&w ©©øÂν׾ -- ©øÂν׾¡@¡@ mU Combined,our results support a role for JH in the nutrition-dependent regulation of extreme mandible growth in this species.
ºî¦X¥H¤W,§Ú̱o¨ìªºµ²ªGÃÒ¹ê¤F¥®ÂζPº¸»X¬O³y¦¨³oª«ºØ²£¥Í·¥¤Ó¤jÃEªºÀç¾i¾¯²K¥[¤W¦³µÛ³o»ò¤@Ó§@¥Î.<x6@ ©©øÂν׾ -- ©øÂν׾¡@¡@ .9'jY In addition, they illuminate mechanisms underlying the evolution of trait proportion, the most salient feature of the evolutionary diversification of the insects.%c ¥t¥~,¥H¤W¤]©úÅãÃÒ©ú¤F(©øÂζPº¸»X)³o¬O¦b©øÂάɸ̳̩úÅã¶i¤Æªº¤ÀÃþ¤W,¯S¼x¤ñ¨Òµo®iªº¤@Óì²z];$ ©©øÂν׾ -- ©øÂν׾¡@¡@ H (Ó¤H»{¬°)²³æªº»¡,´N¬O¥Î¶Pº¸»X¾¯¶qªº²K¥[,Åý©øÂβ£¥Í¤F¤½¥À»P¤jÃE¤j¤pªº®t²§©Êa
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