1/6/11

Strigolactone:  Overview  
!   New  plant  hormone,  which  
were  first  discovered  in  crop  
plants  during  the  infection  
with  the  parasitic  plants  
Striga.  

STRIGOLACTONES   !   Composed  of  tricyclic  lactone  

which  connects  via    an  enol  
ether  bridge  to  
butyrolactone  D  ring  

New  plant  hormone  

Biological  activity  of  strigolactone  

!   Germination  stimulant  
activity  
Other  function?  
q  Increasing  parasite  
infection  
!   The  absence  of  a  natural  plant  parasite  or  a  
!   Host  recognition  signal   mycorrhizal  interaction  for  some  plants  suggested  
for  AMF   strigolactones  have  other  functions.  
q  Enhance  the  
absorption  of  nutrients   !   But,  what  are  these  role(s)?  
from  soil  
  q  Hayward  et  al.  (2009)  à  both  auxin  and  strigolactone  
have  the  capacity  to  modulate  each  other’s  levels  and  
distribution  required  for  axilary  branching  

Shoot  branching     Shoot  branching  

!   Shoot  formation:   !   Mobile  signal  à  strigolactone  or  derivatives  
q  The  initiation  of  axillary  buds   q  ccd7  and  ccd8  mutant  of  rice  and  pea  restore  shoot  
q  The  outgrowth  of  shoot   branching  
q  control  of  their  outgrowth  by  genetically,  hormonally  and   q  Synthetic  strigolactone  (GR24)  can  restore  branching  
environmentally  regulated  signals   inhibition  in  these  mutants  as  well  as  in  max1,  max3  and  
max4  in    Arabidopsis  
!   Previously  only  auxins,  ABA,  and  cytokinins  have  been  
assumed  to  role  shoot  branching   !   In  the  shoot,  strigolactone  signal  transduction  probably  
involves  MAX2  gene,  the  F-­‐box  leucine-­‐rich  repeat  (LRR)  
!   Mutations  in  the  Arabidopsis  MORE  AXILLARY  GROWTH   protein.  
(MAX)  genes  and  orthologous  genes  in  garden  pea  (Pisum   q  Branching  in  max2  plants  is  not  repressed  by  GR24.    
sativum),  rice  (Oryza  sativa)  and  petunia  (Petunia  hybrida)   q  F-­‐box  proteins  function  in  Skp1-­‐Cul1/Cdc53-­‐F-­‐box  (SCF)  E3  
all  lead  to  increased  branching   ubiquitin  ligase  complexes,  which  target  proteins  for  
q  MAX3,  MAX4,  MAX1,  control  the  production  of  an  upwardly   ubiquitination  
mobile  signal  -­‐-­‐-­‐-­‐-­‐-­‐-­‐|  axillary  bud  outgrowth  

1  
 1/6/11  

Shoot  branching    
Auxin    
!   Inhibit  branching  
!   Also  signals  via  SCF  mediated  targeted  protein  
degradation  
q  The  F-­‐box  protein,  TIR1,  and  AFBs  act  as  auxin  receptors  
!   AXR1  à  proper  SCF  function  
q  Mutation  in  AXR1  à  defects  in  downstream  auxin  responses  
q  Appears  to  regulate  SCF  signaling  in  photomorphogenesis  
and  jasmonate  responses  
q  axr1  à  increased  branching,  agravitropic  root  growth,  
!   MAX3  and  MAX4  in  Arabidopsis  encode  carotene   fewer  lateral  root  and  root  hairs,  and  reduced  fertility  due  
dioxygenases  (CCD7  and  CCD8)  à  carotene   to  poor  stamen  elongation  
degradation  into  a  variety  of  product   !   In  Arabidopsis:  
q  Mutation  in  the  MAX  genes  à  increased  stem  conductivity  
!   MAX1,  encodes  a  cytochrome  P450,  is  assumed  to  act   for  auxin  and  increased  expression  of  several  auxin  
downstream  of  the  two  CCDs,  revealed  the  importance   transporter,  including  some  of  the  PIN  family  
 
of  carotene-­‐dioxygenases  (CCDs).  

Interaction  between  auxin  and  

MAX3  and  MAX4  expression  is  reduced  
strigolactone  through  MAX3  and  
by  auxin  depletion  treatments  
MAX4  regulation  
!   A  dose  response  of  MAX3  and  MAX4   !   Decapitation  of  auxin  sources  
expression  to  the  natural  IAA  was  
undertaken  in  the  basal  cauline   !   applying  auxin  transport  
internode  of  intact  axr1-­‐3  and  WT   inhibitor,  NPA  
plants  

! Auxin  positively  regulates  both  

MAX3  (CCD7)  and  MAX4    (CCD8)  
expression  and  this  was  AXR1-­‐
dependent  

Auxin-­‐related  down-­‐regulation  of  

MAX3  and  MAX4  expression  can   bdl-­‐2  
activate  branching  
!   Grafting  studies  revealed  that  WT  roots  can  restore  branching  in  
!   A  semi-­‐dominant  auxin  
max3  and  max4  shoots  to  WT  levels  while  max3  can  not  rescue   response  mutant  that  
max4  shoots  and  vise  versa   confers  defective  
q  MAX3  and  MAX4  are  required  in  the  same  tissue  for  
strigolactone  production  and  their  product  can  move  from  the   primary  root  formation  
root  to  the  shoot  to  inhibit  branching   in  the  embryo  and  
q  Roots  of  axr1-­‐3  à  restore  branching  in  max4  shoots  to  WT   greatly  increased  shoot  
levels,  suggesting  that  axr1-­‐3  roots  are  not  strigolactone  
deficient   branching  in  the  adult  
!   MAX-­‐independent  roles  for  auxin  signaling  in  branching   plant,  particularly  
supression,  but  they  do  not  rule  out  a  role  for  AXR1/TIR1-­‐ obvious  in  
mediated  regulation  of  MAX3  and  MAX4  transcription  in  
modulating  branching  levels.   heterozygous  plant  
q  bodenlos-­‐2  mutant  

2  
 1/6/11  

Graft  transmissibility  of  feedback  signals  

bdl-­‐2  
!   Grafted  of  max2  and  WT  plants  were  
done  to  test  if  feedback  regulation  of  
MAX3  and  MAX4  expression  can  occur  
!   Grafting  experiments   over  long-­‐distances:  
!   Feedback  in  the  shoot  
q  Local  feedback  was  noted  in  the  shoot  
q  For  all  grafts,  the  expression  of  all  genes  
in  the  shoot  was  dependent  only  on  the  
shoot  genotype,  therefore  no  upwardly  
mobile  feedback  signaling  was  detected  

!   Feedback  in  the  root  

q  There  is  a  local  feedback  in  the  root,  but    
downwardly  mobile  feedback  signal  
from  max2  shoots  was  not  able  to  up-­‐
regulated  gene  expression  in  WT  roots  
q  There  is  suggestive  of  some  long  
distance  effect  on  MAX3  and  MAX4  
expression  in  the  rootstock  (depend  on  
shoot  genotype)  

The  mechanisms  of  strigolactone  feedback  

regulation  in  Arabidopsis   Conclusion  
!   Potential  mechanisms  for  the  up-­‐regulation  of  MAX3   ! Auxin-­‐regulation  of  strigolactone  
and  MAX4  gene  expression  in  max  mutants     biosynthetic  genes  in  Arabidopsis  
involves  AXR1/TIR1-­‐regulated  
I.  Direct  effect  of  the  strigolactone  signalling  pathway   Aux/IAA  stability  and  contributes  
II.  Involvement  of  auxin  in  the  feedback  regulation  of   to  branching  inhibition  
MAX3  and  MAX4  
! Auxin  regulation  acts  as  feedback  
III.  Systemic  feedback  regulation  of  MAX3  and  MAX4   mechanism  
!   Increased  auxin  content  in  
condition  of  low  strigolactone  
acting  as  a  downstream  
communicator  to  increase  
strigolactone  biosynthesis  

3