title: Conditional cold avoidance drives between-population variation in germination behaviour in Calluna vulgaris.
abstract: OBJECTIVE
  Across their range, widely distributed species are exposed to a variety of climatic and other environmental conditions, and accordingly may display variation in life history strategies. For seed germination in cold climates, two contrasting responses to variation in winter temperature have been documented: first, an increased ability to germinate at low temperatures (cold tolerance) as winter temperatures decrease, and secondly a reduced ability to germinate at low temperatures (cold avoidance) that concentrates germination towards the warmer parts of the season.
  
  
  METHODS
  Germination responses were tested for Calluna vulgaris, the dominant species of European heathlands, from ten populations collected along broad-scale bioclimatic gradients (latitude, altitude) in Norway, covering a substantial fraction of the species' climatic range. Incubation treatments varied from 10 to 25 °C, and germination performance across populations was analysed in relation to temperature conditions at the seed collection locations.
  
  
  RESULTS
  Seeds from all populations germinated rapidly and to high final percentages under the warmer incubation temperatures. Under low incubation temperatures, cold-climate populations had significantly lower germination rates and percentages than warm-climate populations. While germination rates and percentages also increased with seed mass, seed mass did not vary along the climatic gradients, and therefore did not explain the variation in germination responses.
  
  
  CONCLUSIONS
  Variation in germination responses among Calluna populations was consistent with increased temperature requirements for germination towards colder climates, indicating a cold-avoidance germination strategy conditional on the temperature at the seeds' origin. Along a gradient of increasing temperatures this suggests a shift in selection pressures on germination from climatic adversity (i.e. low temperatures and potential frost risk in early or late season) to competitive performance and better exploitation of the entire growing season.
authors.affiliation: Faculty of Engineering and Science, Sogn og Fjordane University College, PO Box 133, 6851 Sogndal, Norway. joachim.spindelbock@hisf.no
authors.forename: Joachim P
authors.initials: JP
authors.lastname: Spindelböck
authors.forename: Zoë
authors.initials: Z
authors.lastname: Cook
authors.forename: Matthew I
authors.initials: MI
authors.lastname: Daws
authors.forename: Einar
authors.initials: E
authors.lastname: Heegaard
authors.forename: Inger E
authors.initials: IE
authors.lastname: Måren
authors.forename: Vigdis
authors.initials: V
authors.lastname: Vandvik
mesh_terms: D000531:Altitude; D018533:Biomass; D029795:Calluna / Q000502:physiology*; D002980:Climate; D003080:Cold Temperature; D005843:Geography; D018525:Germination / Q000502:physiology*; D009664:Norway; D011891:Rain; D012639:Seeds / Q000502:physiology*
keywords: Calluna vulgaris; cold avoidance; cold stratification; conditional; frost; germination; gradient; heathland; incubation; temperature