title: Genetic variation of piperidine alkaloids in Pinus ponderosa: a common garden study.
abstract: OBJECTIVE
  Previous measurements of conifer alkaloids have revealed significant variation attributable to many sources, environmental and genetic. The present study takes a complementary and intensive, common garden approach to examine genetic variation in Pinus ponderosa var. ponderosa alkaloid production. Additionally, this study investigates the potential trade-off between seedling growth and alkaloid production, and associations between topographic/climatic variables and alkaloid production.
  
  
  METHODS
  Piperidine alkaloids were quantified in foliage of 501 nursery seedlings grown from seed sources in west-central Washington, Oregon and California, roughly covering the western half of the native range of ponderosa pine. A nested mixed model was used to test differences among broad-scale regions and among families within regions. Alkaloid concentrations were regressed on seedling growth measurements to test metabolite allocation theory. Likewise, climate characteristics at the seed sources were also considered as explanatory variables.
  
  
  RESULTS
  Quantitative variation from seedling to seedling was high, and regional variation exceeded variation among families. Regions along the western margin of the species range exhibited the highest alkaloid concentrations, while those further east had relatively low alkaloid levels. Qualitative variation in alkaloid profiles was low. All measures of seedling growth related negatively to alkaloid concentrations on a natural log scale; however, coefficients of determination were low. At best, annual height increment explained 19.4 % of the variation in ln(total alkaloids). Among the climate variables, temperature range showed a negative, linear association that explained 41.8 % of the variation.
  
  
  CONCLUSIONS
  Given the wide geographic scope of the seed sources and the uniformity of resources in the seedlings' environment, observed differences in alkaloid concentrations are evidence for genetic regulation of alkaloid secondary metabolism in ponderosa pine. The theoretical trade-off with seedling growth appeared to be real, however slight. The climate variables provided little evidence for adaptive alkaloid variation, especially within regions.
authors.affiliation: USDA Forest Service, Pacific Northwest Research Station, Corvallis, OR 97331, USA.
authors.forename: Elizabeth A
authors.initials: EA
authors.lastname: Gerson
authors.forename: Rick G
authors.initials: RG
authors.lastname: Kelsey
authors.forename: J Bradley
authors.initials: JB
authors.lastname: St Clair
mesh_terms: D000470:Alkaloids / Q000032:analysis / Q000737:chemistry / Q000235:genetics*; D002980:Climate; D014644:Genetic Variation; D005843:Geography; D016018:Least-Squares Analysis; D016013:Likelihood Functions; D008954:Models, Biological; D019601:Pacific States; D041604:Pinus ponderosa / Q000737:chemistry* / Q000235:genetics* / Q000254:growth & development / Q000378:metabolism; D010880:Piperidines / Q000032:analysis / Q000737:chemistry / Q000378:metabolism*; D010936:Plant Extracts / Q000737:chemistry; D018515:Plant Leaves / Q000737:chemistry; D036226:Seedlings / Q000737:chemistry
chemical_list: D000470:Alkaloids; D010880:Piperidines; D010936:Plant Extracts; C032727:piperidine