Genetics, phosphorus availability, and herbivore-derived induction as sources of phenotypic variation of leaf volatile terpenes in a pine species

Oleoresin produced and stored in pine tree leaves provides direct resistance to herbivores, while leaf volatile terpenes (LVT) in the resin are also powerful airborne infochemicals. Resin concentration and profile show considerable spatial and temporal phenotypic variation within and among pine populations. LVT biochemistry is known to be under genetic control, and although LVT should be plastic to diverse abiotic and biotic environmental factors such as nutrient availability and herbivore attack, little is known about their relative contributions and interactive effects. The aim of this paper was to clarify whether reduced phosphorus availability could increase the LVT concentration and affect the expression of herbivore-derived induced defences, and how plasticity would contribute to the phenotypic variation of LVT. The constitutive and methyl-jasmonate (MeJa) induced LVT concentration and profile were analysed in 17 half-sib Pinus pinaster families growing under two levels of P-availability (complete and P-limited fertilization). Individual terpene concentrations showed large additive genetic variation, which was more pronounced in the control than in MeJa-induced pines. MeJa application did not affect the LVT concentration, but significantly modified the LVT profile by depleting the α-pinene content and reducing the sesquiterpene fraction. Low P-availability strongly reduced plant growth and foliar nutrient concentrations, but did not affect LVT concentration and profile, and did not interact with MeJa-induction. Results indicate a strong homeostasis of LVT concentration to P-availability, and minor changes in the LVT profile due to MeJa-induction. Genetic variation appears to be the main source of phenotypic variation affecting the LVT concentration in this pine species.     

Elevational gradients in plant defences and insect herbivory: recent advances in the field and prospects for future research

Classic research on elevational gradients in plant–herbivore interactions holds that insect herbivore pressure is stronger under warmer, less seasonal climates characteristic of low elevations, and that this in turn selects for increased defence in low‐ (relative to high‐) elevation plants. However, recent work has questioned this paradigm, arguing that it overly simplifies the ecological complexity in which plant–insect herbivore interactions are embedded along elevational gradients. Numerous biotic and abiotic factors vary with elevation, and their simultaneous influences are the focus of current work on elevational gradients in insect herbivory and plant defences. The present review 1) synthesizes current knowledge on elevational gradients in plant–insect herbivore interactions; 2) critically analyses research gaps and highlights recent advances that contribute to filling these gaps; and 3) outlines new research opportunities to uncover underlying mechanisms and build towards a unified theory on elevational gradients. We conclude that the next generation of studies should embrace community complexity – including multi‐trophic dynamics and the multivariate nature of plant defence – and to do so by combining observational data, manipulative experiments and emerging analytical tools.     

Trade‐offs between constitutive and induced defences drive geographical and climatic clines in pine chemical defences

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^o 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.     

Defensive Traits in Young Pine Trees Cluster into Two Divergent Syndromes Related to Early Growth Rate

The combination of defensive traits leads to the evolution of ‘plant defense syndromes’ which should provide better protection against herbivores than individual traits on their own. Defense syndromes can be generally driven by plant phylogeny and/or biotic and abiotic factors. However, we lack a solid understanding of (i) the relative importance of shared evolution vs. convergence due to similar ecological conditions and (ii) the role of induced defense strategies in shaping defense syndromes. We investigate the relative roles of evolutionary and ecological factors shaping the deployment of pine defense syndromes including multiple constitutive and induced chemical defense traits. We performed a greenhouse experiment with seedlings of eighteen species of Pinaceae family, and measured plant growth rate, constitutive chemical defenses and their inducibility. Plant growth rate, but not phylogenetic relatedness, determined the deployment of two divergent syndromes. Slow-growing pine species living in harsh environments where tissue replacement is costly allocated more to constitutive defenses (energetically more costly to produce than induced). In contrast, fast-growing species living in resource-rich habitats had greater inducibility of their defenses, consistent with the theory of constitutive-induced defense trade-offs. This study contributes to a better understanding of evolutionary and ecological factors driving the deployment of defense syndromes.     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

Growth and nutritional response of Pinus pinaster after a large pine weevil (Hylobius abietis) attack

Hylobius abietis is an important pest of coniferous plantations in Europe, to which high mortality, stem deformities, and growth loss are typically attributed. In pine trees, as in other long-lived organisms, there is uncertainty regarding the long-term costs of short-term resistance against invading organisms. We examined the nutritional status of Pinus pinaster after a 2-year long H. abietis attack, measuring needle and phloem N and P concentrations, and the impact of the damage on subsequent growth, survival, and stem deformities over a period of 5 years. The study sites were a P. pinaster family × fertilization trial, and a neighbouring twin trial with similar climate and soil characteristics that was not attacked. Growth losses after the H. abietis attack were important (up to 40%), but restricted to the first years after the attack. Five years after the attack, the annual height increment of pines in the attacked stand was not related to the initial damage suffered, and plants showed regular stems, normal leader dominance, and regular height after 5 years. These findings suggest strong compensatory growth in P. pinaster and indicate relatively high tolerance to the large pine weevil. Needle nutrient concentrations in the healthy stand were, as expected, significantly greater in experimentally fertilized plants, and they were linearly related to those in phloem showing equilibrated stoichiometry both for nitrogen (r = 0.86; P < 0.01; N = 25) and phosphorus (r = 0.84; P < 0.01; N = 25). However, at the attacked stand, nutrient concentrations in the needles did not follow the experimentally manipulated nutrient availability in soils, and phosphorus concentration in the needles was unexpectedly not related to those in the phloem. The pine seedlings attacked by H. abietis showed altered potential of allocating nutrients to their tissues according to the nutrient availability existing in the soil, as well as altered stoichiometry in N and P concentrations among phloem and leaves. Maritime pine seems to be tolerant to the pine weevil attack, at least in the conditions of this study, where pine weevil damage caused a deep alteration of nutrient allocation and nutritional status. Further research is needed to elucidate to what extent altered nutrient allocation may be part of an induced response to the attack or just derived from the vascular injury caused by the weevil wounding in the phloem.