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nitrogenase
N2 NH3
Greensulfur Bacteroides
Prokaryotes Spirochetes
Deinococci
Green, Chlamydiae
nonsulfur
Thermotoga Gram positive
Cyano
bacteria
Rhizobium
Bradyrhizobium
Sinorhizobium
Agrobacterium
Azospirillum
Desulfoivbrio Herbaspirillum
E. coli
Klebsiella
Azotobacter
Proteobacteria
Ralstonia
Burkholderia
Rickettsia
Devosia
Azorhizobium
Sinorhizobium
Methylobacterium
Bradyrhizobium
Proteobacteria Nitrobacter, Afipia
Mesorhizobium,
Aminobacter, Phyllobacterium
Bartonella
Brucella
Sinorhizobium
Ensifer
Rhizobium
Agrobacterium
nitrogenase
N2 + 8 flavodoxin + 8H + 16 MgATP + 18 H2O
- + 2-
GOGAT
glutamate -ketoglutarate + glutamine
Pathway 2
GDH
NH4+ + -ketoglutarate glutamate
Amino acids
proteins
A growing population must eat!
•Estimated that 90% of population will live in tropical and subtropical areas
where (protein-rich) plant sources contribute 80% of total caloric intake.
Consumes 1.4%
of total fossil
fuels annually
rhizobia
Obvious signs of nodulation by common rhizobial species
MEDICAGO LOTUS
(alfalfa) (birdsfoot trefoil)
Pea Plant
R. leguminosarum
nodules
© Simms
Very early events in the Rhizobium-legume symbiosis
chromosome
NodD
nod-gene inducers
from alfalfa roots plasmid
(specificity)
pSym
activated NodD
positively regulates
nod genes
nod genes
Nod factor NodM
biosynthesis
NodC
Nod factor R-group
“decorations”
determine host
specificity
NodB
Nod Factor: a
lipooligosaccharide
Rhizobium Attachment and infection
Nod factor
(specificity)
Invasion through infection tube
Flavonoids
(specificity)
Bacteroid Nitrogen
differentiation fixation
Formation of
nodule primordia
From Hirsch, 1992.
New Phyto. 122, 211-237
• Signals early in infection
– Complex handshaking between legume root
and rhizobium
Correct
signal
Incorrect
signal
Rhizobium encoding GFP from jellyfish as a marker
Infection thread
Enlargement of the
nodule, nitrogen
fixation and
exchange of
nutrients
The Nodulation Process
• Chemical recognition of roots and Rhizobium
• Root hair curling
• Formation of infection thread
• Invasion of roots by Rhizobia
• Cortical cell divisions and formation of nodule
tissue
• Bacteria fix nitrogen which is transferred to
plant cells in exchange for fixed carbon
Inoculation of a mutated Sinorhizobium strain does
not transfer fixed N to the plant
6 days
7 days
wt glnBP5
Genes & Development
11:1194, 1997
Azorhizobium caulinodans
on
Sesbania
Actinorhizal symbioses
Frankia in alder root nodules
Frankia vesicles
Growth response of P. vulgaris plants (45 dpi) inoculated with R. etli strains in
the greenhouse. Images: 1, Noninoculated nonfertilized; 2, inoculated with
CFN42 (wt); 3, inoculated with HP55 (nifHcDK); 4, noninoculated fertilized with
10 mM KNO3–2 mMNH4NO3.
The End