title: Nitrogen limitation and high density responses in rice suggest a role for ethylene under high density stress.
abstract: BACKGROUND
  High density stress, also known as intraspecies competition, causes significant yield losses in a wide variety of crop plants. At the same time, increases in density tolerance through selective breeding and the concomitant ability to plant crops at a higher population density has been one of the most important factors in the development of high yielding modern cultivars.
  
  
  RESULTS
  Physiological changes underlying high density stress were examined in Oryza sativa plants over the course of a life cycle by assessing differences in gene expression and metabolism. Moreover, the nitrogen limitation was examined in parallel with high density stress to gain a better understanding of physiological responses specific to high density stress. While both nitrogen limitation and high density resulted in decreased shoot fresh weight, tiller number, plant height and chlorophyll content, high density stress alone had a greater impact on physiological factors. Decreases in aspartate and glutamate concentration were found in plants grown under both stress conditions; however, high density stress had a more significant effect on the concentration of these amino acids. Global transcriptome analysis revealed a large proportion of genes with altered expression in response to both stresses. The presence of ethylene-associated genes in a majority of density responsive genes was investigated further. Expression of ethylene biosynthesis genes ACC synthase 1, ACC synthase 2 and ACC oxidase 7 were found to be upregulated in plants under high density stress. Plants at high density were also found to up regulate ethylene-associated genes and senescence genes, while cytokinin response and biosynthesis genes were down regulated, consistent with higher ethylene production.
  
  
  CONCLUSIONS
  High density stress has similar but greater impact on plant growth and development compared to nitrogen limitation. Global transcriptome changes implicate ethylene as a volatile signal used to communicate proximity in under dense population growth condition and suggest a role for phytohormones in high density stress response in rice plants.
authors.forename: Maksym
authors.initials: M
authors.lastname: Misyura
authors.forename: David
authors.initials: D
authors.lastname: Guevara
authors.forename: Sanjeena
authors.initials: S
authors.lastname: Subedi
authors.forename: Darryl
authors.initials: D
authors.lastname: Hudson
authors.forename: Paul D
authors.initials: PD
authors.lastname: McNicholas
authors.forename: Joseph
authors.initials: J
authors.lastname: Colasanti
authors.affiliation: Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada. rothstei@uoguelph.ca.
authors.forename: Steven J
authors.initials: SJ
authors.lastname: Rothstein
mesh_terms: D001224:Aspartic Acid / Q000378:metabolism; D005030:Ethylenes / Q000378:metabolism*; D020869:Gene Expression Profiling; D017343:Genes, Plant / Q000235:genetics; D018698:Glutamic Acid / Q000378:metabolism; D055432:Metabolomics; D009584:Nitrogen / Q000378:metabolism*; D012275:Oryza / Q000235:genetics* / Q000254:growth & development / Q000378:metabolism* / Q000502:physiology; D013312:Stress, Physiological
chemical_list: D005030:Ethylenes; D001224:Aspartic Acid; D018698:Glutamic Acid; C036216:ethylene; D009584:Nitrogen