title: Genetic dissection of Al tolerance QTLs in the maize genome by high density SNP scan.
abstract: BACKGROUND
  Aluminum (Al) toxicity is an important limitation to food security in tropical and subtropical regions. High Al saturation on acid soils limits root development, reducing water and nutrient uptake. In addition to naturally occurring acid soils, agricultural practices may decrease soil pH, leading to yield losses due to Al toxicity. Elucidating the genetic and molecular mechanisms underlying maize Al tolerance is expected to accelerate the development of Al-tolerant cultivars.
  
  
  RESULTS
  Five genomic regions were significantly associated with Al tolerance, using 54,455 SNP markers in a recombinant inbred line population derived from Cateto Al237. Candidate genes co-localized with Al tolerance QTLs were further investigated. Near-isogenic lines (NILs) developed for ZmMATE2 were as Al-sensitive as the recurrent line, indicating that this candidate gene was not responsible for the Al tolerance QTL on chromosome 5, qALT5. However, ZmNrat1, a maize homolog to OsNrat1, which encodes an Al(3+) specific transporter previously implicated in rice Al tolerance, was mapped at ~40 Mbp from qALT5. We demonstrate for the first time that ZmNrat1 is preferentially expressed in maize root tips and is up-regulated by Al, similarly to OsNrat1 in rice, suggesting a role of this gene in maize Al tolerance. The strongest-effect QTL was mapped on chromosome 6 (qALT6), within a 0.5 Mbp region where three copies of the Al tolerance gene, ZmMATE1, were found in tandem configuration. qALT6 was shown to increase Al tolerance in maize; the qALT6-NILs carrying three copies of ZmMATE1 exhibited a two-fold increase in Al tolerance, and higher expression of ZmMATE1 compared to the Al sensitive recurrent parent. Interestingly, a new source of Al tolerance via ZmMATE1 was identified in a Brazilian elite line that showed high expression of ZmMATE1 but carries a single copy of ZmMATE1.
  
  
  CONCLUSIONS
  High ZmMATE1 expression, controlled either by three copies of the target gene or by an unknown molecular mechanism, is responsible for Al tolerance mediated by qALT6. As Al tolerant alleles at qALT6 are rare in maize, marker-assisted introgression of this QTL is an important strategy to improve maize adaptation to acid soils worldwide.
authors.affiliation: Nucleus of Applied Biology, Embrapa Maize and Sorghum, Road MG424, km 65, Sete Lagoas, MG 35701-970, Brazil. claudia.guimaraes@embrapa.br.
authors.forename: Claudia T
authors.initials: CT
authors.lastname: Guimaraes
authors.forename: Christiano C
authors.initials: CC
authors.lastname: Simoes
authors.forename: Maria Marta
authors.initials: MM
authors.lastname: Pastina
authors.forename: Lyza G
authors.initials: LG
authors.lastname: Maron
authors.forename: Jurandir V
authors.initials: JV
authors.lastname: Magalhaes
authors.forename: Renato C C
authors.initials: RC
authors.lastname: Vasconcellos
authors.forename: Lauro J M
authors.initials: LJ
authors.lastname: Guimaraes
authors.forename: Ubiraci G P
authors.initials: UG
authors.lastname: Lana
authors.forename: Carlos F S
authors.initials: CF
authors.lastname: Tinoco
authors.forename: Roberto W
authors.initials: RW
authors.lastname: Noda
authors.forename: Silvia N
authors.initials: SN
authors.lastname: Jardim-Belicuas
authors.forename: Leon V
authors.initials: LV
authors.lastname: Kochian
authors.forename: Vera M C
authors.initials: VM
authors.lastname: Alves
authors.forename: Sidney N
authors.initials: SN
authors.lastname: Parentoni
mesh_terms: D000220:Adaptation, Biological / Q000235:genetics*; D000535:Aluminum / Q000633:toxicity*; D001947:Breeding; D002874:Chromosome Mapping; D018628:Gene Dosage; D018506:Gene Expression Regulation, Plant / Q000187:drug effects; D017343:Genes, Plant; D018745:Genome, Plant; D005838:Genotype; D010641:Phenotype; D010802:Phylogeny; D018517:Plant Roots / Q000187:drug effects / Q000235:genetics; D020641:Polymorphism, Single Nucleotide; D040641:Quantitative Trait Loci; D003313:Zea mays / Q000187:drug effects* / Q000235:genetics*
chemical_list: D000535:Aluminum