The influence of physiological traits on climatic niche occupancy and competitive ability in the polyploid plant, Chamerion angustifolium (Onagraceae) Ken A. Thompson, Brian C. Husband, Hafiz Maherali University of Guelph, Department of Integrative Biology Guelph, ON. N1G 2W1 Photo: Brian Husband What is polyploidy? • Whole genome multiplication • Instantaneous “speciation” • Associated with angiosperm diversification (Albert et al. 2013. Science 6165) 2x 4x stemcellthailand.org Amborella trichopoda (wikipedia.org) Polyploidy Influences Plant Function • Often increased cell size in polyploids • Cell size influences organ size – Influences physiology 2x 4x 2x stemcellthailand.org 4x Photo: Brian Husband Polyploidy and Distribution • How do physiological differences between cytotypes influence their geographic distributions? 2x 4x Photo: Brian Husband 1. Do physiological traits influence the climatic niche of diploid and polyploid cytotypes? 2. Do physiological traits influence the competitive abilities of diploid and polyploid cytotypes? Study System: Fireweed • Chamerion angustifolium (Onagraceae) • Diploid (2n = 2x = 36) and tetraploid (2n = 4x = 72) cytotypes • Cytotypes occupy largely distinct geographic ranges Photo: Brian Husband Cytotype Physiology in Fireweed • Tetraploids tolerate drought better than diploids, but are more vulnerable to freezing damage (Maherali et al. 2009. New Phytol. 184) 4x Cytotype Physiology in Fireweed • Tetraploids tolerate drought better than diploids, but are more vulnerable to freezing damage (Maherali et al. 2009. New Phytol. 184) • Caused by differences in xylem conduit diameter 2x 4x 1. Do physiological traits influence the climatic niche of diploid and polyploid cytotypes? 2. Do physiological traits influence the competitive abilities of diploid and polyploid cytotypes? • Tetraploid C. angustifolium tolerate drought – Tetraploid niche will be drier than the diploid niche • Diploid C. angustifolium resist freezing – Diploid niche will be colder than tetraploid niche Characterizing the Climatic Niche • Extracted climate data with GIS – 134 C. angustifolium populations – Pure Diploid, Mixed-Ploidy, Pure Tetraploid • Univariate climate variable comparisons Total Season Season Growing TotalGrowing Water DefificDeficit ( mm )(mm) Moisture Soil Tetraploids occupy driest habitats 100 0 -100 -200 -300 A AB B ANOVA F = 5.732 p < 0.01 Mean Minimum Growing Season Temperature ( °C ) Diploids occupy coldest habitats 8 6 4 2 0 -2 A A B ANOVA F = 31.33 p < 0.001 Physiological traits correlated with climatic niche Do physiological traits influence the climatic niche of diploid and polyploid cytotypes? • Physiological traits generate accurate predictions of cytotype niches in C. angustifolium • Biotic interactions also limit fundamental niche 1. Do physiological traits influence the climatic niche of diploid and polyploid cytotypes? 2. Do physiological traits influence the competitive abilities of diploid and polyploid cytotypes? • Tetraploids can tolerate drought – Tetraploids will be better competitors than diploids in dry soils Experimental Design • 864 plants in 180 pots • 15 competition treatments at 7 densities Applying Irrigation Treatments Aboveground Dry Biomass (g) • Two irrigation treatments • Well watered and drought treatments independently controlled *p < 0.01 ANOVA * * Diploid Tetraploid Analysis of Competition Experiment • Reciprocal-yield model – Multiple linear regression 1 Wš = bš0 + šš Nš + šš” Nš” + Nš Nš” + šš Intracytotypic competition competition Intercytotypic Reciprocal biomass of an individual diploid plant Diploid density Tetraploid density Analysis of Competition Experiment • Reciprocal-yield model – Multiple linear regression 1 Wš = bš0 + šš Nš + šš Nš” + Nš Nš” + šš – Test for difference between šš and šš RECIPROCAL BIOMASS (g) RECIPROCAL BIOMASS (g) DENSITY OF DIPLOIDS DENSITY OF TETRAPLOIDS DENSITY OF DIPLOIDS DENSITY OF TETRAPLOIDS Equal Competitive Abilities Different Competitive Abilities šš = šš” šš ≠ šš” Equal Competitive Abilities in WellWatered Treatment Pots DIPLOID RESPONSE TETRAPLOID RESPONSE šš = šš” šš = šš” Equal Competitive Abilities in Drought Treatment Pots DIPLOID RESPONSE TETRAPLOID RESPONSE šš = šš” šš = šš” Cytotypes are equally competitive Do physiological traits influence the competitive abilities of diploid and polyploid cytotypes? • No difference in competitive ability between cytotypes in either watering treatment • Drought tolerance does not increase tetraploid competitive ability • Physiological traits can be used to predict the climatic niches of C. angustifolium cytotypes • Competition for water probably not restricting cytotype distribution – Drought could influence earlier life stages – Competition and temperature? Thank You! • Acknowledgements – Paul Kron—Flow Cytometry – Mike Mucci—Irrigation System – NSERC Discovery, Canada Foundation for Innovation, Canada Research Chair Program grants to BCH and HM Super Secret Extra Slides http://clipartist.info/clipart/halloween/plain_ghost-1979px.png Ploidy Screening • Used flow cytometry to screen over 300 fruits Number of Nuclei – Compare sample DNA content to a known standard Tomato (DNA Standard) Diploid (2x) Tetraploid (4x) Fluorescence Area Principal Components Analysis • Supports unique climatic niche hypothesis Principal Component 2 3 MIN TEMP MEAN TEMP MAX TEMP MEAN PREC MEAN PET 2 1 0 -1 Pure Diploid Mixed-Ploidy Pure Tetraploid -2 -3 -4 -3 -2 -1 0 1 2 Principal Component 1 3 4 Niche Modelling • Models produce predictions that are similar to the estimated geographic distributions Diploid Niche Modelling • Models produce predictions that are similar to the estimated geographic distributions Tetraploid