Rhizobium and Other N Fixing Bacteria: Nitrogenase

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Rhizobium and other N fixing bacteria

N
2
NH
3

nitrogenase
Ecology of nitrogen-fixing bacteria
N
2
+ 8 flavodoxin
-
+ 8H
+
+ 16 MgATP
2-
+ 18 H
2
O
+ 2OH
-
+ 8 flavodoxin + 16 MgADP
-
+ 16H
2
PO
4
-
+ H
2
nitrogenase
Biological nitrogen fixation:
2NH
4
+
1. Rare, extremely energy consuming conversion
because of stability of triply bonded N
2
2. Produces fixed N which can be directly
assimilated into N containing biomolecules
Ammonia assimilatory cycle:
How nitrogen enters biological pathways
NH
4
+

Amino acids
proteins
Amino acids
proteins
purines
pyrimidines
-ketoglutarate glutamate
GDH
+
glutamate glutamine + ATP + ADP + Pi
GS
+
NH
4
+

glutamine glutamate -ketoglutarate
GOGAT
+
Pathway 1
Pathway 2
N
2
dinitrogen gas
(78% of air)
NH
4
+

ammonium
NO
2
-

nitrite
NO
3
-

nitrate
BIOSPHERE
N
2
O
nitrous oxide
Denitrification
Nitrogen fixation
the Haber Process
and lightning
Denitrification
nitrification
The Nitrogen Cycle
nitrification
N
2
dinitrogen gas
(78% of air)
NH
4
+

ammonium
NO
2
-

nitrite
NO
3
-

nitrate
N
2
O
nitrous oxide
Prokaryotes
Animals
Plants
assimilation
nitrification
uptake
Biological nitrogen fixation
consumption
The Nitrogen Cycle
In 1910 humans consumed 10% of total carbon fixed by photosynthesis, by
2030 it is predicted that 80% will be used by humans.
Estimated that 90% of population will live in tropical and subtropical areas
where (protein-rich) plant sources contribute 80% of total caloric intake.
A growing population must eat!
Combined nitrogen is the most common limiting nutrient in agriculture
Production of nitrogenous fertilizers has plateaued in recent years
because of high costs and pollution

Estimated 90% of applied fertilizers never reach roots and contaminate
groundwater
Consumes 1.4%
of total fossil
fuels annually
Why chemical fertilizers arent the answer
Host plant Bacterial symbiont

Alfalfa Rhizobium meliloti
Clover Rhizobium trifolii
Soybean Bradyrhizobium japonicum
Beans Rhizobium phaseoli
Pea Rhizobium leguminosarum
Sesbania Azorhizobium caulinodans


Both plant and bacterial factors determine specificity
Rhizobium-legume symbioses
legume
rhizobia
Fixed nitrogen
(ammonia)
Fixed carbon
(malate, sucrose)
MEDICAGO
(alfalfa)
LOTUS
(birdsfoot trefoil)
Obvious signs of nodulation by common rhizobial species
Pea Plant
R. leguminosarum
nodules
Pink color is leghaemoglobin a protein
that carries oxygen to the bacteroids
Physiology of a legume nodule
rhizosphere
Flavonoids
nod-gene
inducers
Nod-factor

Very early events in the Rhizobium-legume symbiosis
From Hirsch, 1992.
New Phyto. 122, 211-237
Rhizobium
Formation of
nodule primordia

Bacteroid
differentiation
Nitrogen
fixation
Nod factor
(specificity)
Invasion through infection tube
Attachment and infection
Flavonoids
(specificity)
Enlargement of the
nodule, nitrogen
fixation and
exchange of
nutrients
Nodule development
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
Genes & Development
11:1194, 1997
wild-type mutant
glnB10
glnBP5
6 days
7 days
wt
wt
Genes & Development
11:1194, 1997
Azorhizobium caulinodans
Sesbania
on
Non-symbiotic nitrogen fixation
Cyanobacteria
Anabaena
Nostoc
Aquatic:
Terrestrial and rhizosphere-associated:
Azospirillum
Azotobacter
Acetobacter
Klebsiella
Clostridium
Plant-associated nitrogen fixation:
the endophytes and epiphytes
Acetobacter diazotropicus
Lives as an endophyte of sugarcane and various other monocots and some dicots
On sugarcane
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).
-Co +Co
A nitrogen-fixing fern
Another cyanobacterium on the palm Welfia regia in an
epiphyllic relationship
It is believed that these bacteria transfer some % of fixed N to
the plants through the leaf surfaces
http://ohioline.osu.edu/hyg-fact/3000/3054.html
Symptoms of crown gall
The End

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