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nitrogenase
N2 NH3
Today’s Topics:
Green, Chlamydiae
nonsulfur
Thermotoga Gram positive
Cyano
bacteria
Rhizobium
a Bradyrhizobium
Sinorhizobium
Agrobacterium
b Azospirillum
d
Desulfoivbrio g Herbaspirillum
E. coli
Klebsiella
Azotobacter
Proteobacteria
Ecology of nitrogen-fixing bacteria
Biological nitrogen fixation:
nitrogenase
N2 + 8 flavodoxin- + 8H+ + 16 MgATP2- + 18 H2O
2NH4+ + 2OH- + 8 flavodoxin + 16 MgADP- + 16H2PO4- + H2
+ GS
NH4 + glutamate + ATP glutamine + ADP + Pi
GOGAT
glutamate a-ketoglutarate + glutamine
Pathway 2
GDH
NH4+ + a-ketoglutarate glutamate
Amino acids
proteins
N2
dinitrogen gas
Denitrification (78% of air) Nitrogen fixation
the Haber Process
and lightning
N 2O
nitrous oxide
The Nitrogen Cycle
NH4+
ammonium
Denitrification
BIOSPHERE
NO3-
nitrate
-
nitrification
NO2
nitrification nitrite
The Nitrogen Cycle N2
dinitrogen gas
(78% of air)
N 2O
nitrous oxide nitrification
Prokaryotes
assimilation
NH4+
ammonium
Plants
consumption
Animals
uptake
NO3 -
nitrate
NO2-
nitrite
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
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
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
Non-symbiotic nitrogen fixation
Aquatic:
Cyanobacteria
Anabaena
Nostoc
The aquatic fern Azolla is the only fern that can fix nitrogen. It does
so by virtue of a symbiotic association with a cyanobacterium
(Anabaena azollae).
Another cyanobacterium on the palm Welfia regia in an
epiphyllic relationship
http://ohioline.osu.edu/hyg-fact/3000/3054.html
Agrobacterium tumefaciens “transforms” plant cells
Transgenes
produce
OPINES, unique
amino acid-like
molecules,
as well as plant
hormones
The End
Current approaches to improving biological
nitrogen fixation
and on grapevine
and on grapevine
Malate
to bacteria
nitrogen-
fixing bacteroid
containing Rhizobium
TCA
ATP ADP+Pi
NH4+
to plant
N2 NH4+