L’evoluzione degli orologi biologici

  • 706 views
Uploaded on

L’evoluzione degli orologi biologici - Rudi Costa

L’evoluzione degli orologi biologici - Rudi Costa

More in: Education
  • Full Name Full Name Comment goes here.
    Are you sure you want to
    Your message goes here
    Be the first to comment
    Be the first to like this
No Downloads

Views

Total Views
706
On Slideshare
0
From Embeds
0
Number of Embeds
3

Actions

Shares
Downloads
6
Comments
0
Likes
0

Embeds 0

No embeds

Report content

Flagged as inappropriate Flag as inappropriate
Flag as inappropriate

Select your reason for flagging this presentation as inappropriate.

Cancel
    No notes for slide

Transcript

  • 1. !"#$%&'()%*#+,#-&)+%.%&%-)+/)%&%-)0) 1%,%&2%+3%456 7*)$#.4)58+,)+96,%$6 :)&6*%;+<+=55%/.#+>??@
  • 2. A)%&%-)06&+1BC5BD4
  • 3. E*,.%45#*#+,)+F64%;+GGG+4#0+63
  • 4. G*+HI>@;+6+J.#*0B+645.%*%D#.+*6D#,+K#6*+K60L'#4+,"=.5%'4+,#+:6).6*+,#$)4#,+6+*%MN0&644)0+0).06,)6*+#OP#.)D#*5Q G*5.)-'#,+/C+5B#+,6)&C+%P#*)*-+6*,+0&%4)*-+%2+5B#+&#6$#4+%2+6+B#&)%5.%P#+P&6*5;+,#+:6).6*+,#0),#,+5%+5#45+MB#5B#.+5B)4 /)%&%-)06&+R/#B6$)%'.R+M64+4)DP&C+6+.#4P%*4#+5%+5B#+4'*Q+F%+,%+4%;+B#+0%*2)*#,+6+P&6*5+5%+5B#+,6.SQ+FB#+,6)&C+.BC5BD)0 D%5)%*4+%2+5B#+B#&)%5.%P#T4+&#6$#4+P#.4)45#,+#$#*+)*+5B#+6/4#*0#+%2+4'*&)-B5Q
  • 5. 36.&+$%*+!)**U"4+J&%M#.0&%0S+VHIWXY
  • 6. KZ.-#*+E40B%22+%/4#.$#4+0).06,)6*+.BC5BD4+)*+B'D6*4+)*+/'*S#.+#OP#.)D#*54 (Aschoff, 1969)
  • 7. 1%*+[%*%PS6++6*,+#CD%'.+A#*(#.
  • 8. !"#$%&+D'56*54+N+&%0%D%5%.+605)$)5C+)*+0%*456*5+,6.S*#44;+]]+++++V[%*%PS6+H@IHY ? ++++++++++++>W !"#'+N+>W+B !"#(+^+H@+B !"#)+^+>@+B !"#%*N++___
  • 9. 3&%*)*-+%2+!"#$%&**VH@<WY :)S#+`%'*-;+1%0S#2#&&#. K#22+a6&&+b+:)0B6#&+1%4/64B; A.6*,#)4
  • 10. 9c1+]=:EGd 3N,%D 9E+EeA FB.Nf&C d! 0 1218 !"#) !"#+ Vg6&+++E4PY V#.++++E4*Y !"#, 3!] V45%PY
  • 11. !"#$%&*D1dE+6*,+P.%5#)*+0C0&# ,#&6C D1dE 91=FcGd ? H> >W d#-65)$#+2##,/60S+&%%P+^+,#&6C+6*,+P.%5#)*+456/)&)5C+0.)5)06&
  • 12. 9c1+*#-65)$#+2##,/60S 9c1 !"# 9c1
  • 13. FB#C+6&&+B6$#+0).06,)6*+0&%0S4 3C6*%/605#.)6 E.6/),%P4)4 ].%4%PB)&6 d#'.%4P%.6 a'D6* :%'4#
  • 14. f#*#.)0+D%,#&+2%.+5B#+0).06,)6*+4C45#D G*P'5+5%+5B#+3&%0S 3&%0S+D#0B6*)4D ='5P'5+%2+5B#+3&%0S V2.%D+h)i*#*+6*,+`%'*-;+>??jY
  • 15. 9.%P#.5)#4+%2+0).06,)6*+.BC5BD4 ! 9#.)%,)0)5C+%2+6PP.%O)D65#&C+>W+B%'.4 !+9#.4)45#*0#+)*+0%*456*5+#*$).%*D#*56&+0%*,)5)%*4 !+C*0B.%*)(65)%*+/C+#O5#.*6&+45)D'&) !+F#DP#.65'.#+0%DP#*465)%*+VP#.)%,+,%#4+*%5+0B6*-# M)5B+#O5#.*6&+5#DP#.65'.#k++lH?+mHY
  • 16. ]%+0C6*%/605#.)6+B6$#+6+0&%0S_ N+5B#C+,)$),#+D%.#+5B6*+%*0#+#$#.C+>W+B%'.4 N+*)5.%-#*+2)O65)%*+)4+0).06,)6*+)*+0C6*%/605#.)6 a%M+06*+0).06,)6*+.BC5BD4+)*+0C6*%/605#.)6+/#+$)4'6&)(#,_ N+PB%5%4C*5B#5)0+4C45#D+D)-B5+/#+.#-'&65#,+/C+5B#+0&%0S+Nn ++'4#+%2+5B#+P.%D%5#.+%2+6+-#*#+)*$%&$#,+)*+PB%5%4C4*5B#4)4 D1dE E/'*,6*0#+%2+!(-./+-#*#+D1dE+)4 &#$#& 0).06,)6* >W W< a%'.4+)*+!!
  • 17. g)4'6&)(65)%*+%2+0).06,)6*+.BC5BD4+)*+0C6*%/605#.)6 P+. 9!(-./ 012. 0123 5.#P5%DC0)* 9.%D%5#. !'0)2#.64# +++.#4)456*0#+ f#*%D)0+4#L'#*0#4+2.%D++45"67%6%661(*"0%589:1( hB#*+5B#+P.%D%5#.+)4+605)$65#,;+)*+6+.BC5BD)0+D6**#.;+&'0)2#.64#+D6S#4+/605#.)6+-&%M V[%*,%+#5+6&Q;+9QdQEQQ;H@@oY
  • 18. G,#*5)2)065)%*+%2+0&%0S+-#*#4+)*+0C6*%/605#.)6 :'56-#*#4)4 constant conditions here = constant light (LL) D'56*54+M)5B+*%+ 0).06,)6*+.BC5BD D'56*54+M)5B+*%+ 0).06,)6*+.BC5BD !)/.6.C+%2+M)&,N5CP# +;*"0%589:1(+]dE [%*,%+#5+6&Q+H@@W
  • 19. FB#+<9$.3=+0&'45#.+)4+)DP%.56*5+2%.+0).06,)6*+.BC5BD4 5M%+P.%D%5#.4 kaiA kaiB kaiC &%44+%2+0).06,)6*+.BC5BD+)*+0%*456*5+&)-B5 VG4B)'.6+#5+6&Q;+H@@<Y
  • 20. <9$.3=+64+6+2##,/60S+P.%0#44 kaiA kaiB kaiC KaiA KaiC A64#,+%*+%$#.#OP.#44)%* 45',)#4 KaiB VG4B)'.6+#5+6&Q;+H@@<Y
  • 21. A)%&%-)06&+2'*05)%*+%2+5B#+0).06,)6*+0&%0S hB65+)4+5B#+0).06,)6* 0&%0S+-%%,+2%.+_
  • 22. ++h)&,+5CP#+6*,+D'56*5+45.6)*4+#OB)/)5+,)22#.#*5+2.##+.'**)*-+P#.)%,4 ! p+>o+B ! = 25 h ! = 30 h
  • 23. FB)4+)4+5B#+2).45+.)-%.%'4+,#D%*45.65)%*+)*+6*C+%.-6*)4D+%2+6*+6,$6*56-# 0%*2#..#,+/C+6+0).06,)6*+4C45#D+5%+2)5*#44
  • 24. FB#+q0#*5.6&r+0).06,)6*+0&%0S VD645#.+0&%0SY+)4+&%065#,+)*+5B# /.6)*+6*,+.#&)#4+%*+6/%'5+IX 0&%0S+*#'.%*4+P#.+B#D)4PB#.# 3&%0S+*#'.%*4+6.#+,)$),#,+)*+j -.%'P4+&%065#,+)*+5B#+&65#.6& /.6)*+V&65#.6&+*#'.%*4Y+6*, ,%.46&+/.6)*+V,%.46&+*#'.%*4Y
  • 25. 31` ff FG: FG: 3![ 3`3 9c1 9c1e]AF 9c1e+]AFe+FG:
  • 26. 9B%5%.#0#P5)%*+)*+>#%(%!7$09 9B%5)0+45)D'&)+06*+/# 5.6*4,'0#,+5%+5B#+0&%0S D60B)*#.C+$)6+$)4'6& 6*,e%.+#O5.6%0'&6. 5.6*4,'05)%*+P65BM6C4 Hall,2000
  • 27. :%,#&+2%.+31`+605)%*+)*+>#%(%!7$09 9B%5)0+#*5.6)*D#*5 TIM 6*,+.#4#55)*-+%2 CRY TIM 5B#+0).06,)6*+0&%0S CRY TIM M)5B)*+5B#+/.6)* ,#P#*,4+%*+&)-B5N D#,)65#, CRY TIM ,#-.6,65)%*+%2 PER PER FG:c!c+VFG:Y TIM PER :$? CYC CLK d73!c7 !"# 3`F=9!E: PER
  • 28. :$?+D1dE FG:+P.%5#)* >W H< j H@@jQ++!)-B5+^)*,'0#, ,#-.6,65)%*+%2+FG: H> #OP&6)*4+5B#+2&C+913
  • 29. J.%D+J&)#4+5%+:)0#+6*,+/60S ?
  • 30. 3&%0S+D#0B6*)4D+)*+2&)#4+6*,+D6DD6&4
  • 31. FB#+:$?"0"((*-#*# 1%465%+#5+6&Q;+d'0&#)0+E0),4+1#4#6.0B;+H@@I
  • 32. PBC&%-#*C cav06 gr05 0(@:$? *#)-B/%'.Ni%)*)*-+5.## hu151 cav20 cav04 62 cav14 gr103 w6 61 cav56 gr101 gr15 w11 47 gr13 cav17 gr14 17 hu133 w16 12 cav58 47 cav01 61 w13 gr19 0(@:$?+)4+5B#+*#M+6&&#&#Q+36&)/.65)%* M)5B+%'5-.%'P+4'--#454+6.%4# w14 gr100 0.001 .#0#*5&C+>???NH????+C.4+6-%
  • 33. E+45.'05'.#,+P%&CD%.PB)4D (@:$?+.#P.#4#*54+5B#+6*0#45.6&+6&&#&# 0(@:$?+%.)-)*65#,+)*+%'5B+G56&C+6 2#M+5B%'46*,+C#6.4+6-% ]G1c3FG=dE!+c!c3FG=d AE!Ed3Gdf*c!c3FG=d ]c:=f1E9aG3+cgcdF F6'/#.+":*90;+0)#*0#;+>??I
  • 34. ]).#05)%*6&+4#&#05)%*+65+:$?"0"(( F6i)D6+]+p+NHQjs 0(A(@:$? J'+b+!)+]s+p+NHQ<+*4 F6i)D6+]+p+NHQ@ss F6i)D6+]+p+NHQ>+*4 J'+b+!)+]s+p+NHQW+*4 a[E;+*4 J'+b+!)+]s+p+NoQ@sss HKA**** VF6'/#.+#5+6&;+0)#*0#;+>??IY
  • 35. _
  • 36. *+Vt+6&&#&#4Y !%065)%* !65)5',# A6.0#&%*6 :6&6-6 c60B+2#D6&#+M64+'4#,+5% -#*#.65# )4%2#D6&#+&)*#4 f#*%5CP#,+5B.%'-B 6&&#&#N4P#0)2)0N931
  • 37. P6*)4B*&)*#4 =&,+c'.%P#6**&)*#4 d#M+c'.%P#6**&)*#4
  • 38. #64%*6&)5C >;*?"095%89(:"# F#DP#.65'.# .#P.%,'05)$#+4#64%* 9B%5%P#.)%,
  • 39. !)-B5++]6.S FB#*:$?"0"((++P%&CD%.PB)4D+6*,+31` !NFG: NFG: PKfWNX !#OEN31` 0(@:$?*VHo@<+uHW>H+66Y ATG AGC AGA GTT AGG CAG CTC CAC AAT CAC ATC TGG AAT AAT CAG AAC TTT GAT AAA GTG AAA TCG GTT ATG GAC TGG TTA CTA GCA ACT CCG CAG… s-tim (19 + 1398 aa) ATG AGC AGA GTT AGG CAG CTC CAC AAT CAC ATC TGG AAT AAT CAG AAC TTT ATA AAG TGA AAT CGG TT ATG GAC TGG TTA CTA GCA ACT CCG CAG… !NFG:+V6*,+0(@:$?+2&)#4Yk+.#,'0#,+&)-B5+.#4P%*4)$#*#44 !NFG:+)*5#.6054+M#6S&C+M)5B+31`+"+)*0.#64#,+456/)&)5C 31` FG: 6*,.#&&)+":*90;+0)#*0#;++>??I
  • 40. G*4#054+%$#.M)*5#.)*-+45.65#-C :6.S#,+.#,'05)%*+)*+%$6.)6*+,#$#&%PD#*5+6*, $)5#&&%-#*#4)4 E44%0)65#,+5%+&)2#+B)45%.C+5.6)54+4'0B+64k !)P),+0%*5#*5 1#4)456*0#+5%+45.#44 V0%&,+6*,+456.$65)%*Y !%*-#$)5C
  • 41. 0(@:$?*2&)#4k+B)-B#.+)*0),#*0#+%2+,)6P6'4#+.#-6.,&#44+%2 PB%5%P#.)%, ])6P6'4#+)*0),#*0# 0(@:$? (@:$? a%'.4+%2+&)-B5 &4N5)D+2.#L'#*0C+)4+B)-B#.+)*+5B#+=7Fa+.#2&#05)*-+)54+$#.C .#0#*5+%.)-)* F6'/#.+":*90;+0)#*0#;+>??I
  • 42. E.#+5B#+5)D+#22#054+%*+,)6P6'4#+#O#.5#,+5B.%'-B 5B#+0).06,)6*+0&%0Sv%.+)*,#P#*,#*5&C_ ++31` ++FG: 24 ++9c1 a%.D%*6&+0%*5.%& _ %2+,)6P6'4# 18 6 12 9B%5%P#.)%,)0 0&%0Se0%'*5#. 3).06,)6*+0&%0S
  • 43. :%,)2)#,+2.%D+F6'/#.+#5+6&;+0)#*0#;+>??I
  • 44. ++++++++5)D#&#44 modularrP&#)%5.%PC generP&#)%5.%PC cD#.4%*+#5+6&;+F.#*,4+f#*#5;+>??@