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The apparent failure of invasions by alien pines in Europe has been explained by the co-occurrence of native pine congeners supporting herbivores that might easily recognize the new plants as hosts. Previous studies have reported that exotic pines show reduced tolerance and capacity to induce resistance to those native herbivores. We hypothesize that limited genetic variation in resistance to native herbivores and the existence of evolutionary trade-offs between growth and resistance could represent additional potential constraints on the evolution of invasiveness of exotic pines outside their natural range. In this paper, we examined genetic variation for constitutive and induced chemical defences (measured as non-volatile resin in the stem and total phenolics in the needles) and resistance to two major native generalist herbivores of pines in cafeteria bioassays (the phloem-feeder Hylobius abietis and the defoliator Thaumetopoea pityocampa) using half-sib families drawn from a sample of the population of Pinus radiata introduced to Spain in the mid-19th century. We found (i) significant genetic variation, with moderate-to-high narrow-sense heritabilities for both the production of constitutive non-volatile resin and induced total phenolics, and for constitutive resistance against T. pityocampa in bioassays, (ii) no evolutionary trade-offs between plant resistance and growth traits or between the production of different quantitative chemical defences and (iii) a positive genetic correlation between constitutive resistance to the two studied herbivores. Overall, results of our study indicate that the exotic pine P. radiata has limited genetic constraints on the evolution of resistance against herbivores in its introduced range, suggesting that, at least in terms of interactions with these enemies, this pine species has potential to become invasive in the future. © 2013 Macmillan Publishers Limited All rights reserved 0018-067X/13. Source

Quantitative seed provisioning is an important life-history trait with strong effects on offspring phenotype and fitness. As for any other trait, heritability estimates are vital for understanding its evolutionary dynamics. However, being a trait in between two generations, estimating additive genetic variation of seed provisioning requires complex quantitative genetic approaches for distinguishing between true genetic and environmental maternal effects. Here, using Maritime pine as a long-lived plant model, we quantified additive genetic variation of cone and seed weight (SW) mean and SW within-individual variation. We used a powerful approach combining both half-sib analysis and parent-offspring regression using several common garden tests established in contrasting environments to separate G, E and G × E effects. Both cone weight and SW mean showed significant genetic variation but were also influenced by the maternal environment. Most of the large variation in SW mean was attributable to additive genetic effects (h2=0.55-0.74). SW showed no apparent G × E interaction, particularly when accounting for cone weight covariation, suggesting that the maternal genotypes actively control the SW mean irrespective of the amount of resources allocated to cones. Within-individual variation in SW was low (12%) relative to between-individual variation (88%), and showed no genetic variation but was largely affected by the maternal environment, with greater variation in the less favourable sites for pine growth. In summary, results were very consistent between the parental and the offspring common garden tests, and clearly indicated heritable genetic variation for SW mean but not for within-individual variation in SW.Heredity advance online publication, 27 August 2014; doi:10.1038/hdy.2014.76. Source

Moreira X.,University of California at Irvine | Mooney K.A.,University of California at Irvine | Zas R.,Mision Biologica de Galicia MBG CSIC | Sampedro L.,Research Center Forestal Of Lourizan
Proceedings of the Royal Society B: Biological Sciences

While plant diversity is well known to increase primary productivity, whether these bottom-up effects are enhanced by reciprocal top-down effects from the third trophic level is unknown. We studied whether pine tree species diversity, aphid-tending ants and their interaction determined plant performance and arthropod community structure. Plant diversity had a positive effect on aphids, but only in the presence of mutualistic ants, leading to a threefold greater number of both groups in the tri-specific cultures than in monocultures. Plant diversity increased ant abundance not only by increasing aphid number, but also by increasing ant recruitment per aphid. The positive effect of diversity on ants in turn cascaded down to increase plant performance; diversity increased plant growth (but not biomass), and this effect was stronger in the presence of ants. Consequently, bottom-up effects of diversity within the same genus and guild of plants, and top-down effects from the third trophic level (predatory ants), interactively increased plant performance. © 2012 The Royal Society. Source

Moreira X.,University of California at Irvine | Mooney K.A.,University of California at Irvine | Rasmann S.,University of California at Irvine | Petry W.K.,University of California at Irvine | And 3 more authors.
Ecology Letters

There is increasing evidence that geographic and climatic clines drive the patterns of plant defence allocation and defensive strategies. We quantified early growth rate and both constitutive and inducible chemical defences of 18 Pinaceae species in a common greenhouse environment and assessed their defensive allocation with respect to each species' range across climatic gradients spanning 31o latitude and 2300 m elevation. Constitutive defences traded-off with induced defences, and these defensive strategies were associated with growth rate such that slow-growing species invested more in constitutive defence, whereas fast-growing species invested more in inducible defence. The position of each pine species along this trade-off axis was in turn associated with geography; moving poleward and to higher elevations, growth rate and inducible defences decreased, while constitutive defence increased. These geographic patterns in plant defence were most strongly associated with variation in temperature. Climatic and geographical clines thus act as drivers of defence profiles by mediating the constraints imposed by trade-offs, and this dynamic underlays global patterns of defence allocation. © 2014 John Wiley & Sons Ltd/CNRS. Source

Cendan C.,Mision Biologica de Galicia MBG CSIC | Zas R.,Mision Biologica de Galicia MBG CSIC
Environmental and Experimental Botany

Optimizing the germination timing is crucial for the establishment of new generations. We hypothesized that environmental maternal effects may be relevant in the fine tuning of this trait in a long-lived Mediterranean model tree. We analyzed the influence of maternal genotype, maternal environment and their interaction on the germination success and germination phenology of 8725 Pinus pinaster seeds collected from genotypes clonally replicated in two contrasting environments. Besides maternal genetic effects, the maternal environment significantly affected both the percentage and the timing of germination. Seeds from the more favourable environment germinated 7.5 days earlier and showed higher germination rate (0.93. ±. 0.01 vs 0.85. ±. 0.03). Seed weight significantly influenced germination time, but seed weight differences between maternal environments were not enough to explain this form of transgenerational plasticity. The effect of the maternal environment varied depending on the genotype, indicating that genetic variation in the sensitivity to the maternal environment in this pine species does exist. © 2011 Elsevier B.V. Source

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