Moving nitrogen to the centre of plant defence against pathogens.
BACKGROUND
Plants require nitrogen (N) for growth, development and defence against abiotic and biotic stresses. The extensive use of artificial N fertilizers has played an important role in the Green Revolution. N assimilation can involve a reductase series ( NO3- → NO2- → NH4+ ) followed by transamination to form amino acids. Given its widespread use, the agricultural impact of N nutrition on disease development has been extensively examined.


METHODS
When a pathogen first comes into contact with a host, it is usually nutrient starved such that rapid assimilation of host nutrients is essential for successful pathogenesis. Equally, the host may reallocate its nutrients to defence responses or away from the site of attempted infection. Exogenous application of N fertilizer can, therefore, shift the balance in favour of the host or pathogen. In line with this, increasing N has been reported either to increase or to decrease plant resistance to pathogens, which reflects differences in the infection strategies of discrete pathogens. Beyond considering only N content, the use of NO3- or NH4+ fertilizers affects the outcome of plant-pathogen interactions. NO3- feeding augments hypersensitive response- (HR) mediated resistance, while ammonium nutrition can compromise defence. Metabolically, NO3- enhances production of polyamines such as spermine and spermidine, which are established defence signals, with NH4+ nutrition leading to increased γ-aminobutyric acid (GABA) levels which may be a nutrient source for the pathogen. Within the defensive N economy, the roles of nitric oxide must also be considered. This is mostly generated from NO2- by nitrate reductase and is elicited by both pathogen-associated microbial patterns and gene-for-gene-mediated defences. Nitric oxide (NO) production and associated defences are therefore NO3- dependent and are compromised by NH4+ .


CONCLUSIONS
This review demonstrates how N content and form plays an essential role in defensive primary and secondary metabolism and NO-mediated events.
Institute of Environmental and Rural Science, Aberystwyth University, Edward Llwyd Building, Aberystwyth SY23 3DA, UK.
Luis A J
LAJ
Mur
Institute of Environmental and Rural Science, Aberystwyth University, Edward Llwyd Building, Aberystwyth SY23 3DA, UK.
Catherine
C
Simpson
National Institute of Plant Genome Research, Aruna Asaf Ali Marg, 110067, New Delhi.
Aprajita
A
Kumari
National Institute of Plant Genome Research, Aruna Asaf Ali Marg, 110067, New Delhi.
Alok Kumar
AK
Gupta
National Institute of Plant Genome Research, Aruna Asaf Ali Marg, 110067, New Delhi.
Kapuganti Jagadis
KJ
Gupta
D005308:Fertilizers; D054884:Host-Pathogen Interactions; D009584:Nitrogen / Q000008:administration & dosage / Q000032:analysis* / Q000378:metabolism*; D017672:Nitrogen Compounds / Q000008:administration & dosage / Q000378:metabolism*; D010935:Plant Diseases / Q000382:microbiology*; D010944:Plants / Q000378:metabolism*
D005308:Fertilizers; D017672:Nitrogen Compounds; D009584:Nitrogen
Nitric oxide; Pseudomonas; ammonium; nitrate; nitrate reductase; plant defence; polyamines