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 31° 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.

Summary1. Production of antiherbivore chemical defences is generally assumed to be costly in terms of fitness, although some studies have failed to detect such costs. A convincing explanation is that the expression of fitness costs depends on environmental conditions such as nutrient availability. 2. We performed a greenhouse experiment with 33 half-sib families in order to study the phenotypic plasticity of constitutive and methyl jasmonate-induced chemical defences to soil phosphorus (P) availability, the existence of genetic trade-offs (costs) between growth and the production of those defences and the extent to which P availability may modulate the expression of those costs. 3. We measured some proxies of vegetative fitness (primary growth, secondary growth and total biomass), plant reserves (soluble sugars and starch) and the concentration of quantitative chemical defences (diterpene content in the stem, total polyphenolics and condensed tannins in the needles). 4. Phosphorus availability had a considerable effect, both on the allocation of resources to constitutive and induced defences and on the expression of vegetative costs associated with those chemical defences. Constitutive investment in chemical defences was greater under P-limited conditions for all studied traits. Inducibility of foliar phenolic compounds was greater under P-limited conditions, and it was strongly constrained under high P availability. Availability of P did not affect the inducibility of stem diterpenes. 5. All defensive traits showed significant genetic variation, with different levels of genetic control in constitutive and induced modes, and genetic variation in their inducibility. We found significant negative genetic correlations (i.e. trade-offs) between growth and defensive investment, but costs of chemical defences emerged only in P-limited conditions. Vegetative costs of constitutive defences were detected for stem diterpenes but not for needle phenolics, while costs of induced defences were found for leaf phenolics but not for stem diterpenes. 6. Synthesis. Our results indicate that P availability controls the production of chemical defences in this pine species, influencing the resource allocation to constitutive defences, the inducibility of those defences and the emergence of related vegetative costs. Phosphorus availability thus appears as a major driver in the evolution of pine resistance to insects and a potential factor in maintaining genetic variation in defences.

Methyl jasmonate (MeJa) is a plant chemical elicitor which has been used to artificially induce chemical defensive responses and resistance against herbivores in many plant species in recent decades. In this paper, we studied the effect of exogenous MeJa application at different concentrations (0, 5, 50, and 100 mM) on growth, chemical defences and resistance against the large pine weevil Hylobius abietis L in one year old Pinus pinaster Ait. seedlings. We also investigated whether the local application of MeJa on a basal branch would distally elicit defensive responses or growth reductions on the opposite branch and on the upper main stem. Exogenous application of MeJa induced resin accumulation in the stems, and enhanced resistance against H. abietis. The resin content in plants treated with 100 mM was nearly 2-fold greater, and the pine weevil consumed 80% less phloem than in control plants. However, concentration of total phenolics in needles was lower in plants treated with the higher MeJa dose, probably because of a local toxic effect. We did not detect any effect of MeJa application on plant growth traits. In addition, the application of MeJa on the experimental branches did not significantly affect the resin content, nor growth rates of the opposite untreated control branches. However, the application of MeJa on the experimental branches significantly increased the resin content in the stem and total phenolics in the needles in the apical stem section of the seedling. These results suggest that the systemic effect of the MeJa induction is restricted to the most valuable parts of the seedlings, and should be considered in further studies aimed to understand the systemic resistance of conifers.

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Conventional analysis of spatially correlated data in inadequately blocked field genetic trials may give erroneous results that would seriously affect breeding decisions. Forest genetic trials are commonly very large and strongly heterogeneous, so adjustments for microenvironmental heterogeneity become indispensable. This study explores the use of geostatistics to account for the spatial autocorrelation in four Pinus pinaster Ait. progeny trials established on hilly and irregular terrains with a randomized complete block design and large blocks. Data of five different traits assessed at age 8 were adjusted using an iterative method based on semivariograms and kriging, and the effects on estimates of variance components, heritability, and family effects were evaluated in relation to conventional analysis. Almost all studied traits showed nonrandom spatial structures. Therefore, after the adjustments for spatial autocorrelation, the block and family × block variance components, which were extremely high in the conventional analysis, almost disappeared. The reduction of the interaction variance was recovered by the family variance component, resulting in higher heritability estimates. The removal of the spatial autocorrelation also affected the estimation of family effects, resulting in important changes in family ranks after the spatial adjustments. Comparison among families was also greatly improved due to higher accuracy of the family effect estimations. The analysis improvement was larger for growth traits, which showed the strongest spatial heterogeneity, but was also evident for other traits such as straightness or number of whorls. The present paper demonstrates how spatial autocorrelation can drastically affect the analysis of forest genetic trials with large blocks. The iterative kriging procedure presented in this paper is a promising tool to account for this spatial heterogeneity.

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.

Although maternal environmental effects are increasingly recognized as an important source of phenotypic variation with relevant impacts in evolutionary processes, their relevance in long-lived plants such as pine trees is largely unknown. Here, we used a powerful sample size and a strong quantitative genetic approach to analyse the sources of variation of early seedling performance and to identify seed mass (SM)-dependent and -independent maternal environmental effects in Maritime pine. We measured SM of 8924 individual seeds collected from 10 genotypes clonally replicated in two environments of contrasting quality (favourable and stressful), and we measured seedling growth rate and biomass allocation to roots and shoots. SM was extremely variable (up to 14-fold) and strongly determined by the maternal environment and the genotype of the mother tree. The favourable maternal environment led to larger cones, larger seeds and reduced SM variability. The maternal environment also determined the offspring phenotype, with seedlings coming from the favourable environment being 35% larger and with greater root/shoot ratio. Transgenerational plasticity appears, thus, to be a relevant source of phenotypic variation in the early performance of this pine species. Seed provisioning explained most of the effect of the maternal environment on seedling total biomass. Environmental maternal effects on seedling biomass allocation were, however, determined through SM-independent mechanisms, suggesting that other epigenetic regulation channels may be involved.

Summary1. Current hypotheses predict contrasting roles for natural enemies in determining the success or failure of plant invasions. Differences in plant-induced resistance and tolerance to native herbivores between native and exotic species might contribute to resolve this controversy. 2. We examined the differences between the native Pinus pinaster and the exotic P. radiata in constitutive resistance, inducibility of chemical defences, realized resistance and tolerance to the large pine weevil Hylobius abietis in NW Spain. In this region, both pine species closely coexist and are threatened by the weevil, a harmful phloem feeder that causes extensive mortality and growth reduction in young pine stands. 3. We performed two in vitro cafeteria bioassays, two induction experiments with direct exposure to the weevil and spraying methyl jasmonate and an exhaustive field study of the genetic variation in tolerance and resistance in forestry genetic trials. 4. The weevil significantly preferred the native to the exotic pine when twigs were offered as cut material in Petri dishes. However, the pattern in the field was the opposite, with greater damage on the exotic. Inducibility of stem oleoresin did not differ between species when elicited by the application of methyl jasmonate. However, after a 72-h experimental exposure to the weevil, stem resin content in the native pine was double that in the exotic pine, suggesting a lower capability of the exotic pine to respond to the insect damage. In the field, family relationships between early damage and several pine fitness correlates revealed a significantly greater tolerance of the native pine to the insect damage. Furthermore, only the native pine showed genetic variation in tolerance to the damage. 5. Synthesis. The preference of the herbivore for the native species was counterbalanced by a lower capability for expressing induced resistance to the weevil and reduced tolerance in the exotic species, resulting in no apparent fitness advantage of the exotic P. radiata over the native P. pinaster. Differences in inducibility by and tolerance to native enemies between exotic and native host congeners emerge as key traits for understanding how native enemies might contribute to preventing an introduced species from becoming invasive.