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SVEUILITE U ZAGREBUPRIRODOSLOVNO-MATEMATIKI FAKULTET
BIOLOKI ODSJEKZAVOD ZA ANIMALNU FIZIOLOGIJU
Seminar
Hormoni kukaca
Studentic! Ln G"i#n$i%K&'e(i)! Fi*i&'&(i) end&+"in&( ,u,tV&dite')i +&'e(i)! i*. /"&0. d". ,c. Ve,n Ben+&i%
I*. /"&0. d". ,c. D&1(&) 2i+i%
Z("e34 5678.
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Saetak
P&*nt& )e d )e u* #i$ni4 end&+"ini ,u,t ('ni "e(u't&"
1et3&'i*1 te inte("i" ,e n)e(&e 0un+ci)e u )edin,tenu c)e'inu. Ku+ci4
u ,tdi)u 'i$in+e *3&( ,&( "i(idn&( e(*&,+e'et 1&(u ",ti ,1&/e"i&di$n& +d &d3cu)u ,&) ,t"i e(*&,+e'et i *1i)en)u)u ( n&i14
e%i14 u /"&ce,u n*n&1 /"e,'$en)e. N+&n *dn)e( 'i$in$+&(
/"e,'$en) +u+c u'*i u 0*u d&"1nci)e u &3'i+u +u+u')ice i* +&)e %e ,e
/"&ce,&1 1et1&"0&*e "*iti &d",' )edin+. G'nu u'&(u u "e(u'ci)i
",t i "*&) te "*1nn) i1 u/"& end&+"ini ,u,t +u+c +&)i
,e ,,t&)i &d neu"&,e+"et&"ni9 ,tnic u 1&*(u4 c&"/&"e c"dice4 c&"/&"e
''te4 /"&t&"+'ne #'i)e*de i i*&'i"ni9 end&+"ini9 ,tnic u (n('i)i1 i/"&3i'u4 i*'u$u)u neu"&,e+"et&"ne 9&"1&ne4 /"&t&"+&t"&/ni 9&"1&n4
)ueni'ne 9&"1&ne4 e+di,te"&ide te ne+e /e/tide.
Endokrina regulacija
End&+"in "e(u'ci) u #i&tu +u+c ,e te1e')i n e+di,te"&idi1
:e+di,&n4 56-9id"&+,ie+di,&n;4 )ueni'n&1 9&"1&nu :J
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Z "*'i+u &d /"e,'$en) +&)e u+')u$u)e ,1& /"&1)ene u e/ide"1i,u4
1et1&"0&*&1 ,e d&(>)u i /"&1)ene u inte",tin'n&1 di)e'u +u+c.
:Nti&n;
Slika 1. Gene"'ni u,t"&) end&+"in&( ,u,t +u+c
Kemija i metabolizam hormona
Protorakotropni hormon )e /"i &t+"ieni 9&"1&n +u+c. T& )e
/e/tidni 9&"1&n +&)e( 'u$i /&,te"i&"ni di& 1&*( n*n c&"/&" c"dic
+& &d(&&" n n),+e $i13eni+e i ,ti1u'i")u /"&t&"+'ne #'i)e*de +&)e
,inteti*i")u i 'u$e 9&"1&ne /"e,'$en) :e+di,te"&ide; u 9e1&'i10u.
:Ju"en+;
Juvenilni hormoni ,u ("u/ ci+'i$+i9 ,e,+ite"/en&id +&)i
&,i(u")u ",t 'i$in+e ,/"i)e$)u%i 1et1&"0&*u u /"&ce,u n*n&1
/"e,'$en)e. Jueni'ni 9&"1&n i'i neotenin'u$i /" end&+"ini9 #'i)e*d i*
1&*( *ni9 c&"/&" ''t. U* /"e,'$en)e i(" u'&(u i u /"&du+ci)i ))
u #en+i +u+c. Ve%in +u+c i1 ,1& )ueni'ni 9&"1&n ti/ III4 ,1& u
Le/id&/te" )e /"&n>en& $eti"i ti/ t&( 9&"1&n. T) 9&"1&n ,e i* c&"/&"e
''te &t/u=t u 9e1&'i10u te /utu)e d& ci')ni9 t+i4 "*(">u)e ,e
en*i1i1 )ueni'n&1-9&"1&n e,te"*&1 i )ueni'n&1-9&"1&n e/&+,id
9id"&'*&1 +&)e ti1e ,u/"i1i")u n)e(&& d)e'&n)e. :Ju"en+;
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Slika 2. St"u+tu"n 0&"1u' ne&tenin
Ekdison )e ,te"idni /"&9&"1&n +&)i ,e i*'u$u)e i* /"&t&"+'ni9
#'i)e*d ne+&'i+& ,ti n+&n &t/u=tn) /"&t&"+&t"&/n&( 9&"1&n4
n)e(&i1 ,*"i)en)e1 n,t)e 56-9id"&+,ie+di,&n +&)i )e +tini &3'i+
9&"1&n :Ju"en+;. Z)edni$+i1 i1en&1 ,e ,i ti 9&"1&ni n*i)u
9&"1&ni1 /"e,'$en) i'i ekdisteroidima :Gi'3e"t;. Oni d)e'u)u n
,tnice e/ide"1e e*u)u%i ,e n e+di,&n-"ece/t&" n )e*("i i ,ud)e'u)u u
"e(u'ci)i (en ,/eci?$ni9 * /"e,'$en)e :Ju"en+;.
Slika 3. St"u+tu"n 0&"1u' 56-
9id"&+,ie+di,&n
Morfogenetika ekdisona i juvenilnog hormona
Epiderma i stvaranje kutikule
E/ide"1 +u+c ,e ,,t&)i &d )edn&( ,'&) e'e+t"i$+i /&e*ni9
,tnic +&)e /"&i*&de +uti+u'u +&) /"i')e#e n n)i9. I+& ,e ,tnice
e/ide"1e +u+c ,1t")u di0e"enci"ni1 ,tnic1 +&)e ,t")u
+uti+u'u4 &ne 1&(u d')e di0e"enci"ti i ,t"ti "*'i$ite ,/eci)'i*i"ne
,t"u+tu"e /&/ut $e+in) i d'+. Oi,n& & 9&"1&n'n&1 ut)ec)u4 ,tnice
e/ide"1e 1&(u ,t"ti "*'i$ite ",te +uti+u'4 u+')u$u)u%i +uti+u'e 'i$in+e4
+u+u')ice i &d",'&( ,tdi). :Gi'3e"t;
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U /"&ce,u /"e,'$en)4 /"i d&(>) )e /&'i*i)4 =t& &*n$
&d)n)e +uti+u'e &d ,tnic e/ide"1e. E+di*i)'n 1e13"n +&) ,e
,t" &d) ,t"u +uti+u'u &d &ne +&) te+ t"e3 3iti ,t&"en. St")u ,e
i #e'tin&*ni ,e+"eti +&)i ,e ,,t&)e &d /"&tein i 'i/id te n& /&+uti+u'4
,,t')en &d 9itin i /"&tein. Ot/u=teni en*i1i :&t/u=t i9 ,1
+uti+u'; "*(">u)u ,t"u /"&+uti+u'u ++& 3i &n 1&(' 3iti u+'&n)en +&d
/"e,'$en). N+&n /"e,'$en) d&'*i d& ,*"i)en) n&e +uti+u'e
un+",ni1 /&e*in)e1 /"&tein , 9itin&1. T) /"&ce, 3i u3"*n
/&1&%u 9&"1&n burzikona=t& n&di n *+')u$+ d ,tnice e/ide"1e
&d(&")u n 9&"1&n'ne ,ti1u'u,e +&)i "e(u'i")u /"&ce, /"e,'$en).
J&= )edn 9&"1&n4 e+di,&n4 ,ud)e'u)e u inici)ci)i ,t"n) +uti+u'e4
/"i,utn )e &d /&$et+ "*("dn)e ,t"e +uti+u'e. :Gi'3e"t;
I,t"#in) in it"& ,u /&+*' d )e /&t"e3n 76 -@ i 76- M 56-
9id"&+,ie+di,&n :*n i -e+di,&n; +"&* &d"e>eni /e"i&d "e1en ++& 3i
d&='& d& ,t"n) +uti+u'e. P"&9&"1&n4 e+di,&n :C-e+di,&n; i*'u$u)u
/"&t&"+'ne #'i)e*de i i1 7 +tin&,ti 56-9id"&+,ie+di,&n n ,tnice
e/ide"1e. I+& e+di,&n ne1 ut)ec) n ,tnice e/ide"1e4 #n )e u "'&
"ni1 0*1 /"e,'$en) i /&e% 1et3&'i*1 ,tnic e/ide"1e.Ut)e$e n DNA ,inte*u4 /&)u 9"/&( end&/'*1t,+&( "eti+u'u1 i
/&e%n)e 3"&) 1it&9&nd"i). I,t /&) ,e u&$ i +&d ni,+i9
+&ncent"ci) 56-9id"&+,ie+di,&n. :Gi'3e"t;
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Slika 4. Ve'i$in &d",'e )edin+e , &3*i"&1 n /"i,utn&,t )ueni'n&(
9&"1&n
Sklerotizacija i tamnjenje kutikule
N+&n /"e,'$en) n&&,t&"en +uti+u' &t"dne i $e,t& /&t1ni.
T) /"&ce, ,e *&e ,+'e"&ti*ci). N ,+'e"&ti*ci)u n)$e=%e ut)e$u
neu"&,e+"et&"ni 9&"1&ni4 'i ,u i e+di,te"&idi u+')u$eni.
U 1u94 56-9id"&+,ie+di,&n ut)e$e n ,tnice e/ide"1e i /&)$
t"n,+"i/ci)u (en * d&/ +"3&+,i'*u4 +')u$n&( en*i1 u t1n)en)u.
K,ni)e4 e+di,&n u*"&+u)e &t/u=tn)e neu"&,e+"et&"n&( 9&"1&n4
+u+u')i$in&( 0+t&" * t1n)en)e :&d en('. Pu/"iu1 tnnin( 0ct&"4 PTF; u'i10u +&)i inici" /"&ce, t1n)en).
U e%ine &,t'i9 in,e+t4 t1n)en)e n&e +uti+u'e )e "e(u'i"n&
d"u(i1 neu"&,e+"et&"ni1 9&"1&n&14 3u"*i+&n&1. Ot/u=t ,e i* neu"&n u
3d&1in'ni1 (n('i)i1 u 9e1&'i10u &d19 n+&n 1et1&"0&*e u
&d",'u )edin+u i u*"&+u)e /',ti$n&,t +"i' =t& &1&(u%u)e n)i9&& /&t/un&
=i"en)e te )e /"%en& n)i9&i1 t1n)en)e1. :Gi'3e"t;
Stanino reprogramiranje epiderme
K&d Le/id&/te" i &'e&/te"4 1et1&"0&* */&$in)e u d +&"+!
/d +&ncent"ci)e )ueni'n&( 9&"1&n =t& )e /"%en& n('i1 /&",t&1
+&ncent"ci)e e+di,&n u di)e 0*e. P" 0* &t/u=tn) inici"
1et1&"0&*u u,1)e"n)e1 ,tnic e/ide"1e i* di0e"enci)ci)e u '"u udi0e"enci)ci)u u +u+u')icu. D"u(&4 e%e &t/u=tn)e e+di,&n /&ti$e ,inte*u
+u+u')i$ine +uti+u'e. Jueni'ni 9&"1&n 1e *u,titi 1et1&"0&*u ,1&
+& ne1 56-9id"&+,ie+di,&n. U+&'i+& )e 56-9id"&+,ie+di,&n /"i,utn4
ne1 )ueni'n&( 9&"1&n4 ,tnice e/ide"1e /&,t)u i1une n )ueni'ni
9&"1&n i d&'*i d& ,t"n)e +u+u')i$ine +uti+u'e. A+& )e u* 56-
9id"&+,ie+di,&n /"i,utn i )ueni'ni 9&"1&n4 ,tnice i d')e 1&(u ,t"ti
'i$in$+u +uti+u'u. R*'&( t&1e )e =t& e+di,&n ut)e$e n /"&1)enu
e+,/"e,i)e (en +&)i &1&(u%u)u ,inte*u n&e +uti+u'e. :Gi'3e"t;
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!maginalni diskovi
I1(in'ni di,+&i ,u ,+u/in "e'tin& nedi0e"enci"ni9 ,tnic
/"i,utni9 u 9&'&1et3&'i$+i9 'i$in+i +&)i ",tu ti)e+& ,tdi) 'i$in+e d&+ ,e
1et1&"0&*&1 /"e&3'i+u)u u "*'i$ite ,t"u+tu"e /&/ut &$i)u4 +"i'4 n&(u4(enit'i) i d"u(i9. N)i9& ",t )e +&ntinui"n ti)e+&1 ",t 'i$in+e4
*u,t') ,e u3"*& n+&n /&$et+ "*&) +u+u')ice. N ",t i1(in'ni9
di,+& /&*itin& ut)e$e )ueni'ni 9&"1&n4 d&+ d& di0e"enci)ci)e d&'*i u
/"i,utn&,ti 56-9id"&+,ie+di,&n i &d,utn&,ti )ueni'n&( 9&"1&n =t&
&1&(u%u)e /"i)e'* u ,tdi) +u+u')ice. Ni1e4 56-9id"&+,ie+di,&n /&ti$e
,inte*u DNA4 RNA i /"&tein u di,+&i1 +&)i idu u 1et1&"0&*u4 d&+
)ueni'ni 9&"1&n d)e'u)e nt(&ni,ti$+i. Budu%i d d&'*i d& ,inte*e"*'i$iti9 /"&tein &i,n& & +&13inci)i d)e'&n) & d)u 9&"1&n4
i1/'ici" d )ueni'ni 9&"1&n ne d)e'u)e ,1& in9i3it&"n& e% d u,1)e"
,tni$ne /"&ce,e u di,+&i1. :Gi'3e"t;
Slika 5. P"i,utn&,t /&)edin&( 9&"1&n %e &d'u$iti u +&)u 0*u %e
)edin+ u%i
"lijezde slinovnice i napuhavanje kromosoma
E+di,&n ut)e$e n ,/eci?$n& n/u9in)e /&'iteni9 +"&1&,&1i1 u
#'i)e*d1 ,'in&nic1 d&+"i'c. U *"=n&1 +&"+u #i&t 'i$in+e4
#'i)e*de ,'in&nice ,inteti*i")u ('i+&/"&tein +&)i ,'u#i +& ')e/i'& +&)e %e
?+,i"ti ,t"u +u 'i$in+e * /&d'&(u i td %e te #'i)e*de /"&/,ti. U/"&
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t& n/u9in)e +"&1&,&1 %e &1&(u%iti t"n,+"i/ci)u t&( /"&tein-')e/i'4
n/u9in)e %e ne,tti +d "*in e+di,&n /&",te.
56-9id"&+,ie+di,&n di"e+tn& ut)e$e n "n& n/u9in)e i ,t"n)e
/"&tein +&)i %e ,u/"i1i"ti "n& n/u9in)e i induci"ti +,n&n/u9in)e. Zti14 u "e1en,+&1 /e"i&du &d ,t4 56-9id"&+,ie+di,&n
%e ,e i*(u3iti ++& 3i ,e uti='& te (ene. D+'e4 "e1en,+ /&)n&,t t&(
9&"1&n "e(u'i" n&"1'n& n/u9in)e +"&1&,&1. Jueni'ni 9&"1&n
ne1 &/#en&( u$in+ n te /"&ce,e4 &,i1 =t& in9i3i" +,ni)e 0*e +d
ne1 56-9id"&+,ie+di,&n. P"&teini +&)i n,t)u i1)u /"ete#it& "e(u't&"nu
0un+ci)u u d')n)i1 0*1 "*&). :Gi'3e"t;
Slika6. N/u9n)e +"&1&,&1
Masno tkivo
M,n& t+i& +u+c )e e+i'entn& )et"i i 1,n&1 t+iu +"')e=n)+.
Ti)e+&1 'i$in$+&( #i&t &n& "*(">u)e une=ene nut"i)ente i u /"i/"e1i *
1et1&"0&*u /&9"n)u)e 1te"i)'e +&)i ne ",tu. U ne+i9 "&d& 'i$in$+&
1,n& t+i& ,inteti*i" i &t/u=t ne+&'i+& e'i+i9 /"&tein :866 +D; +&)i ,e
n+u/'))u u 9e1&'i1?. Ti /"&teini ,e n+u/'))u u ("nu'1 i "%)u u
1,n& t+i& +d ,e *3&( 1et1&"0&*e 'i$in+ /"e,tne 9"niti. K&d ne+i9
",t d&'*i d& n+u/')n) ti9 /"&tein *3&( ,1n)en) +&ncent"ci)e
)ueni'n&( 9&"1&n4 d&+ ,e u D. melanogaster/&n&n& "%n)e /"&tein
u 1,n& t+i& i ,t"n)e ("nu' "e(u'i" /&e%n)e1 +&ncent"ci)e 56-
9id"&+,ie+di,&n. 56-9id"&+,ie+di,&n u t&) 0*i t+&>e" /&ti$e ,inte*u ne+i9
n&i9 /"&tein u 1,n&1 t+iu. Ti)e+&1 t")n) 1et1&"0&*e4 u 1,n&1
t+iu ,e /&)')u)u ut&0(ne +u&'e *3&( /"i,utn&,ti 56-9id"&+,ie+di,&n4
n+&n =t& ,e ,1n)i +&ncent"ci) )ueni'n&( 9&"1&n. :Gi'3e"t;
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Hormonalna kontrola reprodukcije
Z "*'i+u &d +"')e=n)+4 +u+ci i1)u 9&"1&ne +&)i u/"'))u
",t&1 i "*&)e1 ++& 3i +&nt"&'i"'i "e/"&du+ci)u u &d",'&1 ,tdi)u. U
&d",'&1 ,tdi)u corpora allata /&n&n& */&$in)e , ,e+"eci)&1
)ueni'n&( 9&"1&n +&)i u/"') &&(ene*&1 u #en+i te )e +&d ne+i9 #n
* ,*"i)en)e ,/&'ni9 #'i)e*d i "e/"&du+tin& /&n=n)e +&d 1u#)+.
P"&t&"+'n #'i)e*d e%in&1 de(ene"i" ti)e+&1 "*&) u &d",'u
)edin+u4 i*'u$in)e e+di,&n u 9e1&'i10u i'i ))nu ,tnicu td &3'))u
))nici +&)i /&,t)u end&+"ini &"(n i ti1e "e(u'i")u &&(ene*u i'i "*&)
e13"i). :Gi'3e"t;
Juvenilni hormon i oogeneza
U'&(e )ueni'n&( 9&"1&n u &&(ene*i ,e 1i)en))u &i,n& & ,tdi)u4
u+')u$u)u ,*"i)en)e /"eite'&(eni9 &&cit4 inici)ci)u ,inte*e ite'&(enin
u 1,n&1 t+iu i indu+ci)u un&, ite'&(enin u &&cite.
Jueni'ni 9&"1&n ,ti1u'i" /"eite'&(eni ",t 0&'i+u' ++& 3i &ni
/&,t'i +&1/etentni /"i1iti ite'&(enin u +,ni)i1 0*1 &&(ene*e.e"4 u 1,n&1 t+iu ,u /"&n>eni
cit&,&'ni "ece/t&"i * )ueni'ni 9&"1&n. J&= )edn u'&( )ueni'n&( 9&"1&n
)e i un&, ite'&(enin u &&cite =t& ,e &$itu)e /&e%n)e1 inte"ce'u'"n&(
/"&,t&"4 'i ,1n)en)e1 ,tni$n&( &'u1en *3&( +tici)e /u1/i +&)e
i*3cu)u te+u%inu i* ,tnice. :Gi'3e"t;
#loga ekdisona u reprodukciji enki
N /"i1)e"u ne+i9 +&1"c )e /"i /ut /&+*n& d e+di,&n i("
#nu u'&(u u "e/"&du+ci)i +u+c. N+&n &3"&+ +"')u4 e+di,&n ,e &t/u=t
i* ))ni+ i /"et" u 56-9id"&+,ie+di,&n +&)i d)e'u)e n 1,n& t+i& i
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u*"&+u)e ,inte*u i &t/u=tn)e ite'&(enin4 /"i1"n& /&)$)u%i
t"n,+"i/ci)u (en * ite'&(enin. K,ni)e )e /&+*n& d ,ud)e'u)e i u
,*"i)en)u ))ni9 ,tnic +&d Drosophile. S1t" ,e d )ueni'ni 9&"1&n
,ti1u'i" t+i& 3d&1en4 &"i)e4 n ,e+"eci)u e+di,&n =t& &nd /&ti$e
/&)$nu ,inte*u ite'&(enin u 1,n&1 t+iu. U "*'i$iti9 ",t +u+c
e+di,&n ,e /"&i*&di u "*'i$iti1 t+ii1. T+& /"i1)e"ice u Orthoptera i
Lepidoptera ))nici ,t")u e+di,&n ne/&,"edn& /"i)e *dn)e 0*e
&&(ene*e4 u d"u(i9 ,e ,inteti*i" u 0&'i+u'"ni1 ,tnic1 i &t/u=t u
))ne ,tnice i inici" ,t"n)e ,e"&*ne +uti+u'e. :Gi'3e"t;
Slika7.S*"i)en)e &&cit +&1"c n+&n 9"n)en) +"')u
$eurosekretorni hormoni
Neu"&,e+"et&"ni 9&"1&ni ,u $e,ti +&d +u+c4 &t/u=t)u ,e +&
&d(&&" n &+&'i=ne ,i(n'e4 3i& n),+e i'i unut"=n)e4 ci') i1 )e
&d"#n)e 9&1e&,t*e &"(ni*14 u+')u$u)u%i "e(u'ci)u 1et3&'i*1
u(')i+&9id"t i 'i/id4 9&1e&,t*u &de4 ut)e$u n ,t"u+tu"nu /&)n&,t
+uti+u'e4 /&n=n)e i &t/u=tn)e /"&t&"+&t"&/n&( 9&"1&n +&)i "e(u'i"
+tin&,t /"&t&"+'ne #'i)e*de. U* #i$ne ,i(n'e4 corpora allata )e
t+&>e" "e(u'i"n neu"&9&"1&n'ni1 0+t&"i1.
%dipokinetki hormon &%KH'
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T) #n neu"&,e+"et&"ni /"&tein ,ud)e'u)e u 1et3&'i*1u
u(')i+&9id"t i 'i/id. Kd n+&n 8-76 1inut 'et +&ncent"ci) t"e9'&*e
:=e%e" u 9e1&'i1? +u+c; ,e ,1n)i4 i,/u=t ,e AK< i* ("ndu'"n&(
"e#n) corpore cardiace te 1&3i'i*i" u,+'di=tene "e*e"e t"i('ice"id i*
1,n&( t+i +&)e 1i=i%i &nd +&"i,te u &3'i+u di('ice"id.
Me9ni*1 d)e'&n) di/&+ineti$+&( 9&"1&n )e /"e+& cAMP-.
P&e%n)e1 +&ncent"ci)e cAMP- +tii" ,e cAMP-&i,n /"&tein +in*
+&) +tii" t"i('ice"id 'i/*u4 ,'i$n& d)e'&n)u d"en'in i ('u+(&n +&d
,i,c. T+&>e"4 /&+*n& )e d AK< d)e'u)e n +"nitin-ceti'-
t"n,0e"*u4 1it&9&nd"i),+i en*i1 +&)i ,ti1u'i" 'i/idni 1et3&'i*1 i un&,
ci'ni9 ("u/ u 1it&9&nd"i)e.
Slika 8. St"u+tu" AK