????? (post-translational regulation)

1
???????(?5? 11/10)??????????
2
??
??????
??????
???? (translational regulation)
?????
????? ?????
????? (post-translational regulation)
???
3
??????????????????????
??!
4
???(?????)????
???????????????????????????????????????
???????????????????????????????
????(??????)????????????????????????????
??????????????????????????????
???????????? ? ????????????? ? ???????????????(??)
?????????????
5
?????????????????????
? ????????????????
6
????????(TIR1)????? Nature 446, 621-622 (2007)
?????? Nature 435, 441-445 (2005)
benzoic acid
l-tryptophan
1-NAA
2,4-D
IAA
???(???)?3H-IAA ? SCFTIR1??????
?? ?????????TIR/AFB??????????? F-box????? TIR1,
AFB1, AFB2, AFB3 ????(?? Proceedligs of the
National Academy of Science (PNAS) 10622540
(2009))
7
??????????????
????? ??????(??)

• ??????????? Protein Kinase
• ??????????????
• ???/?????????(?????)
• ?????????????????
• ??????????????????
• ??????
• ????????(C?????CDPK)
• cAMP???(A????)
• ????????(MAP????)

8
???????????????Two-Component Signal
Transduction Pathways in Arabidopsis. Hwang et
al. Plant Physiology, 2002, 129500515
Figure 1.   Schematic representation of the
two-component and the multistep phosphorelay
signaling systems. A, The prototypical
two-component pathway uses a single phosphoryl
transfer event between a His protein kinase and
its cognate response regulator. B, The multistep
His-to-Asp phosphorelay system in which a
His-containing phosphotransfer protein serves as
a phosphoryl acceptor and donor between the
hybrid protein kinase and the response regulator.
In yeast (Saccharomyces cerevisiae) osmosensing,
the phosphorelay connects to an MAPK cascade
(Wurgler-Murphy and Saito, 1997). In Arabidopsis
cytokinin signaling, a response regulator
directly regulates its target gene expression
(Hwang and Sheen, 2001). The vertical bars
represent transmembrane domains. H, His D, Asp
P, phosphoryl group.
9

• ????????????????????????
• Hypothetical scheme for the pathway for
  low-temperature signal transduction in
  Synechocystis Suzuki et al. Biochem. Soc. Trans.
  (2000) 28, 628630

???????????
???????? ? HiK33???????????
HiK33????? ??????????
10
??????????????/?????? ??????????(??)

• ??(??????)?????????(X)??????
• ????????(??)
• X????????????????
• X????????????(??)?????????????????(mimic)

???? (?????)
???? (????)
A B C D
11
??????????????????????????? Five Histidine
Kinases Perceive Osmotic Stress and Regulate
Distinct Sets of Genes in Synechocystis
Paithoonrangsarid et al. J. Biol. Chem.,
(2004) 279 53078-53086
12
?????????
Figure 9.   Model of the two-component signal
transduction pathways in Arabidopsis. The
cytokinin signal is perceived by multiple His
protein kinases at the plasma membrane. Upon
perception of the cytokinin signal, His protein
kinases initiate a signaling cascade via the
phosphorelay that results in the nuclear
translocation of AHPs (Hwang and Sheen, 2001).
Activated AHPs may interact with sequestered ARRs
or ARR complexes, transfer the phosphate to the
receiver domain of its cognate B-type ARR,
releasing these activation-type ARRs from
putative repressors in the nucleus. The
dephosphorylated AHP shuttles back to the
cytosol, where it can be rephosphorylated. The
liberated ARRs bind to multiple cis elements in
the promoter of target genes. The activation of
the repressor-type ARRs as primary cytokinin
response genes provides a negative feedback
mechanism. In addition to the CTR1 signaling
pathway, additional ethylene signaling pathways
could be mediated by two-component components
(Lohrmann and Harter, 2002). Red light and
cytokinin signaling is converged at ARR4. ARR4
stabilizes the active form of PHYB by inhibiting
dark reversion (Sweere et al., 2001). Stress and
Glc may also modulate two-component signaling
(Urao et al., 1998 F. Rolland and J. Sheen,
unpublished data). RD, Response domain BD, DNA
binding domain AD, transactivation domain PM,
plasma membrane N, nucleus R, putative
repressor FR, far-red light
13
??? MAPKinase?
Nature 451, 789-795 (2008) "Dual control of
nuclear EIN3 by bifurcate MAPK cascades in C2H4
signalling." Yoo et al.
Model of the bifurcate MAPK cascades in ethylene
signalling. The two EIN3 phosphorylation sites
(P) with opposite functions are marked. Without
ethylene, CTR1 directly or indirectly inactivates
MKK9MPK3/6 and probably activates downstream
MAPKs to phosphorylate T592 to promote EIN3
degradation. Ethylene inactivates CTR1 for
MKK9MPK3/6 activation and T174 phosphorylation
to stabilize EIN3. Arrow and blunt ends indicate
positive and negative regulations, respectively.
ACS, ACC synthase MKKK, MAPKKK C, cytoplasm N,
nucleus.
(?????????????HP??)

• ?????????

?? ?? ????
14
??????????DREB?(???????????)
?? ?
??
???? ?
???? ?
15
????????????????????? ???????2007?5????
?????? (??????????)
16
??????????????????????

• ?????
• ?????
• (?????????)
• Trends in plant science, 3411-412 (1998)??

CAM???????CDPK???????PKPK???????????PP?????
???????CNA/B?????????A/B(CBLCNB-like)?CAM???(C
a???)
17
???CBL?????

• CBLs/CIPKs
• CBL19 (Calucinulin-B like proteins)
• CIPK125 (CBL-interacting protein kinase)

Black CIPKs BlueCBL1 RedCBL9
2, 11-17, 19-25
3, 5, 10
1, 4, 6, 8,18
7, 9
Drought-responsive
Cold-responsive
Sheng Luan et al.
18
?????????????Two-hybrid?
19
The CBLCIPK network in plant calcium Signaling.
Sheng Luan Trends in Plant Science Volume 14,
Issue 1, 37-42 (2008)
20
??????????????
?????????????
???????????????
Aquaporins and water homeostasis in
plants Kjellbom et al. TIPS, 1999, 4308
(????????)

• (CDPK and Aquaporins)

????????Ca?????????
21
?????????/??????
Ca???????(?????) ?????
????????? (??-2????-3??)
?????????????
22
??????GFP(2008???????? ????)
Ca2??????(??)?GFP(??)
??????? 2009?11????
???????????? (????)
23
????/?????????????
24
?????????????????????? (????????????????)