Variation of early reproductive allocation in multi-site genetic trials of Maritime pine and Aleppo pine

Luis Santos, Eduardo Notivol, Rafael Zas, Maria Regina Chambel, Juan Majada, José Climent

Abstract


Life histories in Mediterranean pines are well known to be closely related to different fire and disturbance regimes. Variation in these factors is also reflected in reproductive strategies at the specific and intraspecific level. Specifically, the onset of reproduction is a crucial stage for any organism because it has profound implications on fitness. In this paper we focused on the intraspecific variation and plasticity in the threshold size for reproduction and reproductive allocation in two Mediterranean pines, assessed at the onset of reproduction, when trade-offs between reproduction and growth are expected to be greater. Replicated common garden provenance and progeny trials of Aleppo pine and Maritime pine were used to estimate genetic parameters for reproduction and vegetative growth, as well as variation in plasticity in reproductive strategies at the intraspecific level. In both species, high variation among populations was found for both threshold size for reproduction and for reproductive allocation. Reproductive allocation was also highly variable within populations and showed moderate to high values of heritability and high coefficients of additive genetic variation. These results indicate a high genetic control of reproductive traits, while high additive genetic variation is maintained, allowing to face selective pressures. Moreover, reproductive strategies although plastic, showed low genotype x environment interaction, and intraspecific variation was highly consistent across trial sites both at the population and the family levels. The former data confirm the strong genetic control of reproductive traits in these species. Finally, the fact that Aleppo pine starts its reproductive phase as female while Maritime pine can start reproducing either as male or female deserves further attention.

Keywords


Pinus halepensis; Pinus pinaster; genetic variability; plasticity; trade-offs; early reproductive strategies; threshold size for reproduction

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References


Agee J.K., 1998. Fire and pine ecosystems. In: Ecology and biogeography of Pinus (Richardson D.M., ed). Cambridge University Press, Cambridge, UK. pp. 193-218.

Bonser S.P., Aarssen L.W., 2001. Allometry and plasticity of meristem allocation throughout development in Arabidopsis thaliana. Journal of Ecology 89, 72-79. http://dx.doi.org/10.1046/j.1365-2745.2001.00516.x

Brett M.T., 2004. When is a correlation between non-independent variables «spurious»? Oikos 105, 647-656. http://dx.doi.org/10.1111/j.0030-1299.2004.12777.x

Bucci G., González-Martínez S.C., Le Provost G., Plomion C., Ribeiro M.M., Sebastiani F., Alia R.,Vendramin G.G., 2007. Range-wide phylogeography and gene zones in Pinus pinaster Ait. Revealed by chloroplast microsatellite markers. Molecular Ecology 16, 2137-2153. http://dx.doi.org/10.1111/j.1365-294X.2007.03275.x PMid:17498237

Climent J., Prada M.A., Calama R., Chambel M. R., De Ron D.S., Alia R., 2008. To grow or to seed: Ecotypic variation in reproductive allocation and cone production by young female Aleppo pine (Pinus halepensis, Pinaceae). American Journal of Botany 95, 833-842. http://dx.doi.org/10.3732/ajb.2007354 PMid:21632409

Cobb N.S., Trotter R.T., Whitham, T.G., 2002. Longterm sexual allocation in herbivore resistant and susceptible pinyon pine (Pinus edulis). Oecologia 130, 78-87.

Gil L., López R., García-Mateos Á., Gonzálezdoncel I., 2009. Seed provenance and fire-related reproductive traits of Pinus pinaster in central Spain. International Journal of Wildland Fire 18, 1003-1009. http://dx.doi.org/10.1071/WF08101

Glazier D.S., 2002. Resource-allocation rules and the heritability of traits. Evolution 56, 1696-1700. PMid:12353762

González-Ochoa A.I., López-Serrano F.R., De Las Heras J., 2004. Does post-fire forest management increase tree growth and cone production in Pinus halepensis? Forest Ecology and Management 188, 235-247. http://dx.doi.org/10.1016/j.foreco.2003.07.015

Grivet D., Sebastiani F., González-Martínez S.C., Vendramin G.G., 2009. Patterns of polymorphism resulting from long-range colonization in the Mediterranean conifer Aleppo pine. New Phytologist 184, 1016-1028. http://dx.doi.org/10.1111/j.1469-8137.2009.03015.x PMid:19732348

Kang K.S.,Bila A.D., Harju A.M., Lindgren D., 2003. Estimation of fertility variation in forest tree populations. Forestry 76, 329-344. http://dx.doi.org/10.1093/forestry/76.3.329

Karlsson P.S., Méndez M., 2005. The resource economy of plant reproduction. In: Reproductive allocation in plants. Elsevier, Amsterdam, The Netherlands. pp. 1-40. http://dx.doi.org/10.1016/B978-012088386-8/50001-6

Keeley J.E., Zedler P.H., 1998. Evolution of life histories in Pinus. In: Ecology and biogeography of Pinus. (Richardson D.M., ed). Cambridge University Press, Cambridge, UK. pp. 193-218.

Koenig W.D., Knops J.M.H., Carmen W.J., Stanback M.T., Mumme R.L., 1994. Estimating acorn crops using visual surveys. Canadian Journal of Forest Research 24, 2105-2112. http://dx.doi.org/10.1139/x94-270

Koenig W.D., Knops, J.M.H., 2000. Patterns of annual seed production by northern hemisphere trees: a global perspective. American Naturalist 155, 59-69. http://dx.doi.org/10.1086/303302 PMid:10657177

Kozlowski J., 1992. Optimal allocation of resources to growth and reproduction: implications for age and size at maturity. Trends in Ecology & Evolution 7, 15-19. http://dx.doi.org/10.1016/0169-5347(92)90192-E

Martíns P., Sampedro L., Moreira X., Zas R., 2009. Nutritional status and genetic variation in the response to nutrient availability in Pinus pinaster. A multisite field study in Northwest Spain. Forest Ecology and Management 258, 1429-1436. http://dx.doi.org/10.1016/j.foreco.2009.06.041

Matziris D., 1997. Variation in growth, flowering and cone production in a clonal seed orchard of Aleppo pine grown in Greece. Silvae Genetica 46, 224-228.

Méndez M., Karlsson P.S., 2004. Between-population variation in size-dependent reproduction and reproductive allocation in Pinguicula vulgaris (Lentibulariaceae) and its environmental correlates. Oikos 104, 59-70. http://dx.doi.org/10.1111/j.0030-1299.2004.12335.x

Merilä J., Sheldon B.C., 2000. Genetic architecture of fitness and nonfitness traits: empirical patterns and development of ideas. Heredity 83, 103-109. http://dx.doi.org/10.1046/j.1365-2540.1999.00585.x

Mueller R., Wade B., Gehring C., Whitham T., 2005. Chronic herbivory negatively impacts cone and seed production, seed quality and seedling growth of susceptible pinyon pines. Oecologia 143, 558-565. http://dx.doi.org/10.1007/s00442-005-0029-0 PMid:15800748

Ne'eman G., Goubitz S., Nathan R., 2004. Reproductive traits of Pinus halepensis in the light of fire – a critical review. Plant Ecology 171, 69-79. http://dx.doi.org/10.1023/B:VEGE.0000029380.04821.99

Richardson D.M. (ed), 1998. Ecology and biogeography of Pinus. Cambridge University Press, Cambridge, UK.

Richardson D.M., Rundel P.W., Jackson S.T., Teskey R.O., Aronson J., Bytnerowicz A., Wingfield M.J., Proches S., 2007. Human impacts in pine forests: past, present, and future. Annual Review of Ecology Evolution and Systematics 38, 275-297. http://dx.doi.org/10.1146/annurev.ecolsys.38.091206.095650

Roff D.A., 2000. Trade-offs between growth and reproduction: an analysis of the quantitative genetic evidence. Journal of Evolutionary Biology 13, 434-445. http://dx.doi.org/10.1046/j.1420-9101.2000.00186.x

Rutter M.T., Fenster C.B., 2007. Testing for Adaptation to Climate in Arabidopsis thaliana: a calibrated common garden approach. Ann Bot 99, 529-536. http://dx.doi.org/10.1093/aob/mcl282 PMid:17293351 PMCid:2802957

Sampedro L., Moreira X., Martíns P., Zas R., 2009. Growth and nutritional response of Pinus pinaster after a large pine weevil (Hylobius abietis) attack. Trees-Structure and Function 23, 1189-1197. http://dx.doi.org/10.1007/s00468-009-0358-4

Schlichting C.D., 2002. Phenotypic plasticity in plants. Plant Species Biology 17, 85-88. http://dx.doi.org/10.1046/j.1442-1984.2002.00083.x

Schmida A., Lev-Yadum S., Goubittz S., Ne'eman G., 2000. Sexual allocation and gender segregation in Pinus halepensis, P. brutia and P. pinea. In: Ecology, biogeography and management of Pinus halepensis and P. brutia forest ecosystems in the Mediterranean Basin. (Ne'eman G., Trabaud L., eds). Backhuys Publishers, Leiden, The Netherlands. pp. 91-104.

Schmidtling R.C., 1981. The inheritance of precocity and its relationship with growth in Loblolly pines. Silvae Genetica 30, 188-192.

Seifert T., Müller-Starck G., 2009. Impacts of fructification on biomass production and correlated genetic effects in Norway spruce (Picea abies L. Karst.). European Journal of Forest Research 128, 155-169. http://dx.doi.org/10.1007/s10342-008-0219-5

Sgro C.M., Hoffmann A.A., (2004). Genetic correlations, tradeoffs and environmental variation. Heredity 93, 241-248. http://dx.doi.org/10.1038/sj.hdy.6800532 PMid:15280897

Sultan S.E., 2003. Phenotypic plasticity in plants: a case study in ecological development. Evolution & Development 5, 25-33. http://dx.doi.org/10.1046/j.1525-142X.2003.03005.x PMid:12492406

Tapias R., Climent J., Pardos J.A., Gil L., 2004. Life histories of Mediterranean pines. Plant Ecology 171, 53-68. http://dx.doi.org/10.1023/B:VEGE.0000029383.72609.f0

Thompson I., Mackey B., Mcnulty S., Mosseler A., 2009. Forest resilience, biodiversity, and climate change. A synthesis of the biodiversity/resilience/stability relationship in forest ecosystems. Secretariat of the Convention on Biological Diversity, Montreal, Canada. Technical Series no. 43. 67 pp.

Wesselingh R.A., Klinkhamer P.G.L., De Jong T.J., Boorman L.A., 1997. Threshold size for flowering in different habitats: effects of size-dependent growth and survival. Ecology 78 2118-2132.

Yasumura Y., Hikosaka K., Hirose T., 2006. Resource allocation to vegetative and reproductive growth in relation to mast seeding in Fagus crenata. Forest Ecology and Management 229, 228-233. http://dx.doi.org/10.1016/j.foreco.2006.04.003

Zas R., Sampedro L., Prada E., Fernándezlópez J., 2005. Genetic variation of Pinus pinaster Ait. seedlings in susceptibility to the pine weevil Hylobius abietis L. Ann For Sci 62, 681-688. http://dx.doi.org/10.1051/forest:2005064




DOI: 10.5424/fs/2010193-9109

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