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Local adaptation versus inbreeding depression in marginal populations of a Mediterranean alpine plant: are they worthy
of conservation in a context of climate change?
Morente-López, J., García-Fernández, A., Lara-Romero, C., Rubio-Teso, M.L., Ruiz, R., Sánchez, A., Iriondo, J.M.
Área de Biodiversidad y Conservación, Universidad Rey Juan Carlos, E-28933 Móstoles, Madrid, Spain.
• Genetically impoverished populations
• Inbreeding depression• Maladaptation
• Not necessarily depauperate for variation in ecologically relevant traits.
• Locally adapted
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations:• grow under suboptimal environmental conditions• great fluctuations and high probability of extinction
Soulé (1973)
?
Kawecki, T. J. 2008. Annu. Rev. Ecol. Evol. Syst. 39:321–342.
Soulé M. 1973. Annu. Rev. Ecol. Sys. 4:165-187Lande R. 1994. Evolution 48: 1460–1469.Whitlock MC. 2003. Genetics 164: 767–779.
Lande (1994), Whitlock (2003) Kawecki (2008)
Mediterranean alpine environments: highly vulnerable to global warming
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations
Central populations
temperature rainfall
Experimental gene flow between populations:• assessment of inbreeding depression and geneflow of
adaptive/maladaptive value• management tool to assist marginal populations
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations
Central populations
central – marginal geneflow:• Genetic diversity (Holt & Gomulkiewicz, 1997)• Maladaptive alleles or gene combinations
(Kirkpatrick & Barton, 1997)
Holt RD, Gomulkiewicz R. 1997. Am Nat 149:563-572Kirkpatrick M, Barton NH. 1997. Am Nat 150:1-23
Experimental gene flow between populations:• assessment of inbreeding depression and geneflow of
adaptive/maladaptive value• management tool to assist marginal populations
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations
Central populations
marginal-marginal geneflow• Genetic diversity & adaptive alleles or gene combinations (Sexton et al. 2011)
Sexton JP, Strauss SY, Rice KJ. 2011. PNAS 108:11704-11709
Aim: • To assess whether marginal populations at the lowest elevation of
Mediterranean alpine plants are locally adapted/maladapted to the environmental conditions that will prevail with global warming
Introduction Aims Materials & Methods Results & Discussion Conclusions
• Circum-mediterranean alpine chamaephyte• Central System at the lowest latitude of the
distibution range:• Sierra de Béjar• Sierra de Gredos• Sierra de Guadarrama
• Elevation range: 1900 – 2500 m
Introduction Aims Materials & Methods Results & Discussion Conclusions
Silene ciliata Pourret
Introduction Aims Materials & Methods Results & Discussion Conclusions
• Central population
• Marginal populations
GuadarramaGredosBéjar
Central vs. marginal populations
(Giménez-Benavides et al. 2011)
Giménez-Benavides, L., Albert, M.J., Iriondo, J.M. & Escudero, A. Ecography (2011) 34:85-93
Introduction Aims Materials & Methods Results & Discussion Conclusions
Central population from same mountain range
Marginal population
Marginal population from same mountain range
X 6 marginal populations
F1F2
F3
Experimental design:• Seeds obtained in common garden conditions from
artificial crossings simulating different types of geneflow
Experimental design:
• Ex situ sowing experiment at optimum conditions in growth chamber and greenhouse environment (mother plant x type of cross x block x pop. = 8640 seeds).
• In situ sowing experiment at the locations of the 6 marginal populations (mother plant x type of cross x block x pop. = 24000 seeds)
Introduction Aims Materials & Methods Results & Discussion Conclusions
Introduction Aims Materials & Methods Results & Discussion Conclusions
Ex situ germination experiment
F1<F2, F3
F1=F2, F3
Germination = geneflowi + populationj + (geneflow x population)ij + motherplantk(j)
Introduction Aims Materials & Methods Results & Discussion Conclusions
Ex situ germination experiment
RUI AGI CAM SES NAJMORF1<F2,F3F1<F2,F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression No inbreeding depression
F1<F2,F3
Inbreeding depression
F1F2
F3
GuadarramaGredosBéjar
Introduction Aims Materials & Methods Results & Discussion Conclusions
In situ germination experiment
F1<F3 F1>F2
F1=F2,F3 F1<F2
Germination = geneflowi + populationj + (geneflow x population)ij + motherplantk(j)
Introduction Aims Materials & Methods Results & Discussion Conclusions
In situ germination experiment
RUI AGI CAM SES NAJMORF1>F2F1<F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression
No inbreeding depression
F1<F2
Adaptation from MOR
F1F2
F3
Maladaptation from RUI
GuadarramaGredosBéjar
Introduction Aims Materials & Methods Results & Discussion Conclusions
In situ germination experiment
RUI AGI CAM SES NAJMOR
Local adaptation (Giménez-Benavides et al., 2007)Inbreeding depression (García-Fernández et al., 2012)
GuadarramaGredosBéjar
Giménez-Benavides L, Escudero A, Iriondo JM. 2007. Ann Bot 99:723-734.García-Fernández A, Iriondo JM, Escudero A.2012. Oikos 121: 1435-1445.
Introduction Aims Materials & Methods Results & Discussion Conclusions
RUI AGI CAM SES NAJMOR
F1F2
F3
F1>F2F1<F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression
No inbreeding depression
F1<F2
Adaptation from MOR
Maladaptation from RUI
In situ
F1<F2,F3F1<F2,F3 F1=F2,F3 F1=F2,F3
Inbreeding depression No inbreeding depression
F1<F2,F3
Inbreeding depression
Ex situ F1=F2,F3
Introduction Aims Materials & Methods Results & Discussion Conclusions
RUI AGI CAM SES NAJMOR
F1F2
F3
F1>F2F1<F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression
No inbreeding depression
F1<F2
Adaptation from MOR
Maladaptation from RUI
In situ
F1<F2,F3F1<F2,F3 F1=F2,F3 F1=F2,F3
Inbreeding depression No inbreeding depression
F1<F2,F3
Inbreeding depression
Ex situ F1=F2,F3
He= 0.63FIS= 0.33Ne= 10
He= 0.75FIS= 0.32Ne= 25
He= 0.66FIS= 0.19Ne= 12
He= 0.70FIS= -0.12Ne= inf
He= 0.70FIS= 0.39Ne= 66
He= 0.51FIS= 0.31Ne= 131
Introduction Aims Materials & Methods Results & Discussion Conclusions
RUI AGI CAM SES NAJMOR
F1F2
F3
F1>F2F1<F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression
No inbreeding depression
F1<F2
Adaptation from MOR
Maladaptation from RUI
In situ
F1<F2,F3F1<F2,F3 F1=F2,F3 F1=F2,F3
Inbreeding depression No inbreeding depression
F1<F2,F3
Inbreeding depression
Ex situ F1=F2,F3
He= 0.63FIS= 0.33Ne= 10
He= 0.75FIS= 0.32Ne= 25
He= 0.66FIS= 0.19Ne= 12
He= 0.70FIS= -0.12Ne= inf
He= 0.70FIS= 0.39Ne= 66
He= 0.51FIS= 0.31Ne= 131
Introduction Aims Materials & Methods Results & Discussion Conclusions
RUI AGI CAM SES NAJMOR
F1F2
F3
F1>F2F1<F3 F1=F2,F3 F1=F2,F3F1=F2,F3
Inbreeding depression
No inbreeding depression
F1<F2
Adaptation from MOR
Maladaptation from RUI
In situ
F1<F2,F3F1<F2,F3 F1=F2,F3 F1=F2,F3
Inbreeding depression No inbreeding depression
F1<F2,F3
Inbreeding depression
Ex situ F1=F2,F3
He= 0.63FIS= 0.33Ne= 10
He= 0.75FIS= 0.32Ne= 25
He= 0.66FIS= 0.19Ne= 12
He= 0.70FIS= -0.12Ne= inf
He= 0.70FIS= 0.39Ne= 66
He= 0.51FIS= 0.31Ne= 131
Leimu R , Fischer M. 2008. PLoS ONE 3: e4010
Introduction Aims Materials & Methods Results & Discussion Conclusions
RUI AGI CAM SES NAJMOR
F1F2
F3
• Seedling survival and size (up to July): No differences between geneflow treatments
• On-going experiment
In situ
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations
Central populations
F1F2
F3
Little evidence of adaptive geneflow between marginal populations: • Mimulus laciniatus (Sexton et al., 2011)
Sexton JP, Strauss SY, Rice KJ. 2011. PNAS 108:11704-11709
Introduction Aims Materials & Methods Results & Discussion Conclusions
• Some evidence of central populations providing genetic diversity that reduces inbreeding depression (Holt & Gomulkiewicz, 1997)
• No evidence of maladaptive alleles or gene combinations from central populations (Kirkpatrick & Barton, 1997)
Marginal populations
Central populations
F1F2
F3
Holt RD, Gomulkiewicz R. 1997. Am Nat 149:563-572Kirkpatrick M, Barton NH. 1997. Am Nat 150:1-23
Introduction Aims Materials & Methods Results & Discussion Conclusions
Marginal populations
Central populations
F1F2
F3
Genetic differentiation in functional traits between central and marginal populations:• Tolerance to water stress (García-Fernández et al. 2013).• Flowering phenology: S. ciliata (unpublished results), Armeria caespitosa (Lara-
Romero et al., 2014)
García-Fernandez A, Iriondo JM, Bartels D, Escudero A.2013 Plant Biol 15:93-100.Lara-Romero C, García-Camacho R, Escudero A, Iriondo JM. 2014. Bot J Linn Soc 176:384-395.
Introduction Aims Materials & Methods Results & Discussion Conclusions
Conclusions
1. High incidence of inbreeding depression in marginal populations.
2. Both adaptive/maladaptive geneflow between marginal populations at the germination stage.
3. No maladaptive geneflow between central and marginal populations at the germination stage.
4. Variation of responses to geneflow depending on the marginal population are probably associated to differences in effective population size, genetic diversity and population inbreeding.
5. Response to geneflow in a given marginal population is also dependent on temporal variation of environmental conditions
Local adaptation versus inbreeding depression in marginal populations of a Mediterranean alpine plant: are they
worthy of conservation in a context of climate change?
Acknowledgements:
• C. Diaz, G. Escribano, S. Prieto, P. Tabares, S. Eleazar, L. Cano & L. Martinez • Parque Nacional de la Sierra de Guadarrama• Parque Regional de la Sierra de Gredos• Reserva de la Biosfera de la Sierras de Béjar y Francia• AdAptA Project CGL2012-33528, Spanish National R&D&I Plan
