title: Intra-plant variation in nectar sugar composition in two Aquilegia species (Ranunculaceae): contrasting patterns under field and glasshouse conditions.
abstract: OBJECTIVE
  Intra-specific variation in nectar chemistry under natural conditions has been only rarely explored, yet it is an essential aspect of our understanding of how pollinator-mediated selection might act on nectar traits. This paper examines intra-specific variation in nectar sugar composition in field and glasshouse plants of the bumblebee-pollinated perennial herbs Aquilegia vulgaris subsp. vulgaris and Aquilegia pyrenaica subsp. cazorlensis (Ranunculaceae). The aims of the study are to assess the generality of extreme intra-plant variation in nectar sugar composition recently reported for other species in the field, and gaining insight on the possible mechanisms involved.
  
  
  METHODS
  The proportions of glucose, fructose and sucrose in single-nectary nectar samples collected from field and glasshouse plants were determined using high performance liquid chromatography. A hierarchical variance partition was used to dissect total variance into components due to variation among plants, flowers within plants, and nectaries within flowers.
  
  
  RESULTS
  Nectar of the two species was mostly sucrose-dominated, but composition varied widely in the field, ranging from sucrose-only to fructose-dominated. Most intra-specific variance was due to differences among nectaries of the same flower, and flowers of the same plant. The high intra-plant variation in sugar composition exhibited by field plants vanished in the glasshouse, where nectar composition emerged as a remarkably constant feature across plants, flowers and nectaries.
  
  
  CONCLUSIONS
  In addition to corroborating the results of previous studies documenting extreme intra-plant variation in nectar sugar composition in the field, this study suggests that such variation may ultimately be caused by biotic factors operating on the nectar in the field but not in the glasshouse. Pollinator visitation and pollinator-borne yeasts are suggested as likely causal agents.
authors.affiliation: Estación Biológica de Doñana, Consejo Superior de Investigaciones Científicas, Avenida de María Luisa s/n, E-41013, Sevilla, Spain. azucanto@tunku.uady.mx
authors.forename: Azucena
authors.initials: A
authors.lastname: Canto
authors.forename: Ricardo
authors.initials: R
authors.lastname: Pérez
authors.forename: Mónica
authors.initials: M
authors.lastname: Medrano
authors.forename: María Clara
authors.initials: MC
authors.lastname: Castellanos
authors.forename: Carlos M
authors.initials: CM
authors.lastname: Herrera
mesh_terms: D031884:Aquilegia / Q000378:metabolism*; D050260:Carbohydrate Metabolism; D035264:Flowers / Q000378:metabolism; D005632:Fructose / Q000378:metabolism; D005947:Glucose / Q000378:metabolism; D013045:Species Specificity; D013395:Sucrose / Q000378:metabolism
chemical_list: D005632:Fructose; D013395:Sucrose; D005947:Glucose