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Juvenile hormone regulates extreme mandible growth in male stag beetles.s0?`
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Gotoh H, Cornette R, Koshikawa S, Okada Y, Lavine LC, Emlen DJ, Miura T.-v~\C
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Laboratory of Ecological Genetics, Graduate School of Environmental Science, Hokkaido University, Sapporo, Japan.HWc
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The morphological diversity of insects is one of the most striking phenomena in biology.-{6/
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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.<%
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Remarkable examples of an insect trait with exaggerated proportions are the mandibular weapons of stag beetles.&T
¸û·|Åý¤Hª`·N¨ìªºÄ´¦p¦³µÛ¸Ø±i¤ñ¨Ò©øÂίS¼x.§â¤jÃE·íªZ¾¹ªºÁê§ÎÂÎ;&'0*
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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,bcK
stag beetle mandibles vary widely in size among males, and this variable growth results from differential larval nutrition.34h^`k
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However, the mechanisms responsible for coupling nutrition with growth of stag beetle mandibles (or indeed any insect structure) remain largely unknown.=[
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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.J{
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We then investigate the putative role of JH in the development of the nutritionally-dependent, phenotypically plastic mandibles,@QCH
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.Gc
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Increased JH signaling during the early prepupal period increased the proportional size of body parts, and this was especially pronounced in male mandibles,EU|wb
enhancing the exaggerated size of this trait.H`m
¦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.  {@IT |
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Combined,our results support a role for JH in the nutrition-dependent regulation of extreme mandible growth in this species. ºî¦X¥H¤W,§Ú­Ì±o¨ìªºµ²ªGÃÒ¹ê¤F¥®ÂζPº¸»X¬O³y¦¨³oª«ºØ²£¥Í·¥¤Ó¤jÃEªºÀç¾i¾¯²K¥[¤W¦³µÛ³o»ò¤@­Ó§@¥Î.FE
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In addition, they illuminate mechanisms underlying the evolution of trait proportion, the most salient feature of the evolutionary diversification of the insects.\N794
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