Constitutive Resin Research Articles

Constitutive resin ducts formation in Pinus sylvestris responds to water availability and temperature in southern Siberia

Constitutive resin ducts (RDs) are a distinctive anatomical trait in the xylem of most conifers, playing a clear defensive role against biotic and abiotic factors. Although resin duct dynamics are well documented in Mediterranean environments, less attention has been dedicated to continental climate. In this study, we evaluated the occurrence and climate response of resin ducts in Pinus sylvestris L. trees in five sites along a 3422 km transect of increasing continentality in the forest-steppe of southern Siberia over the 1966–2016 period. Our results suggest a strong climatic dependence of tree growth and resin duct production under the Siberian continental climate conditions. Tree-ring width and RDs abundance were positively affected by summer precipitation and negatively influenced by temperature and solar radiation, suggesting a link between RDs formation and the water deficit period that limits RW. However, the RDs climate signal was generally weaker, with the strongest climate response occurring later in the season, showing independent signals when the RW effect was removed. The site played a key role as an explanatory factor of the climate response variability. Although no clear pattern was found along the transect, potential differences are due to phenotypic variations between the P. sylvestris populations. Warming-induced drought conditions during the growing season, associated to ongoing climate warming, might affect tree growth and resin ducts production with an increase of this critical defensive structure abundance in conifers growing in the forest-steppe in southern Siberia.

Resin Canal Traits Variation in Pinus spp. with Different Susceptibility to the Pine Wood Nematode

Different studies have emphasized the influence of resin canal traits in the susceptibility of pine trees to the pine wood nematode (PWN) Bursaphelenchus xylophilus. Resin canals can facilitate the PWN migration through the stem and are involved in the accumulation of volatile terpenes in the xylem in response to the pathogen, inducing tracheid embolisms. In this work, we conducted a PWN inoculation experiment under greenhouse conditions to investigate the anatomical traits of constitutive resin canals among seven Pinus species with different degrees of susceptibility to the PWN: P. canariensis, P. halepensis, P. pinea and P. taeda were grouped into a ‘low-susceptible group’, and P. pinaster, P. radiata, and P. sylvestris were grouped into a ‘high-susceptible group’. The high-susceptible group presented higher xylem radial growth, wider constitutive canals in the cortex, lower frequency of constitutive canals in the cortex, and smaller constitutive canals in the xylem than the low-susceptible group. The size of constitutive cortical canals was positively related to the number of seedlings colonized by the PWN, suggesting that wider canals facilitated migration. The inoculation of the PWN increased the frequency and diminished the mean area of canals in the xylem, and the high-susceptible group showed more frequency of induced xylem canals than the low-susceptible group. Additionally, the high-susceptible group presented larger radial growths in the xylem than the low-susceptible group. These results suggest a role of resin canal traits on Pinus spp. susceptibility to the PWN. Nevertheless, the high interspecific variability found in these traits within each susceptibility group evidences the importance of other factors in the susceptibility to the PWN.

Predicting resin pockets and blemishes in radiata pine lumber from log properties

Background: Resin pockets and blemishes in pruned logs of radiata pine (Pinus radiata D.Don) can reduce the value of clear and moulding grades of lumber. External resin features (ERF) on the bark of the logs have proved an effective method of predicting the incidence of resin pockets in the lumber. Resin canals have been associated with resin blemishes in radiata pine, and could prove useful in improving the prediction of the grade recovery of lumber. Methods: Pruned butt logs of radiata pine trees from two forests in the North Island, New Zealand, were selected for low, moderate, and severe levels of external resin features (ERF) on the bark, and for low, average, and high resin canal diameter, frequency and brightness from breast height increment cores. The relationships were evaluated between these properties, and the lumber resin features and grade recovery of the logs. Results: The number of resin pockets, the blemish rating, and the percentage of boards with resin streaks and resinous heartwood increased, and the recovery of clears and moulding grade boards and the lumber value declined, with the severity of the ERF class of the logs. Multiple regression models gave good predictions of the grade recovery and loss of lumber value, using the log ERF class, volume, heartwood content, and number of Type 1 resin pockets on the ends of the logs, as independent variables. The resin canal properties did not improve the regression models. Resin blemishes were associated with Type 2 resin pockets, and were more frequent in the forest where false growth rings were present. This suggests the constitutive resin flow from resin canals, rather than the resin canal size and frequency, is more important in determining the incidence of resin blemishes. Conclusions: The prediction of the grade recovery of lumber using the ERFs of radiata pine logs, was supplemented by the log volume, heartwood content and number of Type 1 resin pockets on the ends of the logs. The environmental factors that drive the constitutive resin enrichment of resin canals, such as drought conditions that give rise to false growth rings, could be useful in improving the prediction of grade recovery for forest stands.

Growth and defense characteristics of whitebark pine (Pinus albicaulis) and lodgepole pine (Pinus contorta var latifolia) in a high-elevation, disturbance-prone mixed-conifer forest in northwestern Montana, USA

Recent, widespread tree mortality in the western U.S. resulting from changes in climate, pathogens, insect activity, and forest management practices has led to concerns for many ecologically and culturally important species. Within conifers, resin-based defenses have long been recognized as a primary defense mechanism against a variety of insects and pathogens. Oleoresin produced by trees contain complex mixtures of terpenoids that have numerous insecticidal and fungicidal properties. Research has also identified links between resin duct characteristics and increased probability of survival during bark beetle outbreaks. Whitebark pine (Pinus albicaulis) is a culturally significant high elevation species that provides numerous ecological services within subalpine and alpine ecosystems. Whitebark pine has co-evolved with a suite of biotic and abiotic disturbances. Individual trees allocate resources towards growth and resin-based defenses, making it a good candidate species to evaluate growth and defense relationships and tradeoffs. In this study we compared constitutive resin chemistry, tree growth and resin duct anatomy between similarly aged whitebark and lodgepole pine (P. contorta var latifolia) growing in proximity within a disturbance-prone, mixed-conifer forest in northwestern Montana. These two host species have varying degrees of historical exposure to mountain pine beetle. Our research yields four important findings. First, we did not find evidence of a tradeoff between tree growth and tree defenses (resin duct morphology and resin chemistry). This suggests that trees growing under favorable field conditions can experience high growth rates and still allocate ample resources towards defense. Second, we found that resin ducts and constitutive mono- and sesqui- terpenes were not correlated in lodgepole pine while duct production and area were positively related to constitutive monoterpenes, and duct size and area were positively related to constitutive sesquiterpenes, in whitebark pine. The lack of distinct, consistent relationships between these defensive features suggests that both whitebark and lodgepole pine trees present beetles with numerous, complex combinations of resin-based defenses. Third, based on constitutive terpene profiles, bark beetles are more likely to enter lodgepole pine but more likely to successfully elicit mass attacks in whitebark pine, which agrees with beetle attack and success patterns observed in the field. Fourth, overstory competition, particularly by Engelmann spruce (Picea engelmannii), can influence tree defenses, specifically by reducing constitutive terpene concentrations in lodgepole and whitebark pine. Competitive tree interactions could lead to altered bark beetle-conifer interactions as host and nonhost species migrate in response to changing climate. Our results suggest that strategies designed to support whitebark pine populations can benefit from better understanding interactions among growth, competition and physical and chemical defenses in response to multiple disturbance.

Heritability and characteristics of resin ducts in Pinus oocarpa stems in Michoacán, Mexico

Abstract Despite the extensive distribution and use of Pinus oocarpa in Mexico, knowledge on the range of genetic variation and magnitude of quantitative parameters of the defensive structures of the species is rare. Our study attempts to fill this gap by providing a comprehensive assessment of resin duct traits in mother trees and their offsprings in an open-pollinated trial of P. oocarpa. Resin ducts are fundamental structures of the defense mechanisms of the coniferous trees. They vary depending on the age of the tree, the genotype, the species, and may be influenced by environmental factors. We investigated intraspecific variation among families of the resin duct system in secondary vascular tissues (wood and inner bark) of P. oocarpa trees. Our study attempts to explore the variance and the possible genetic control of these defensive structures. We hypothesized that the resin duct features analyzed would vary with the tree’s genetic make-up. We analyzed samples from two groups of trees. First, from mature mother trees selected as superior in resin yield. Second, from their six-year-old off-springs established in a progeny trial. Axial and radial ducts showed differences in their size and quantity, between wood and bark and between ages. The axial duct density showed differences among families, and the heritability estimates were moderate. In P. oocarpa trees selected as phenotypically superior in resin yield, the size of its constitutive resin ducts is a less variable trait, and the differences in axial duct density can be attributed to genetic factors.

Interspecific variation in spruce constitutive and induced defenses in response to a bark beetle-fungal symbiont provides insight into traits associated with resistance.

Differences in defensive traits of tree species may predict why some conifers are susceptible to bark beetle-fungal complexes and others are not. A symbiotic fungus (Leptographium abietinum (Peck) M.J. Wingf.) associated with the tree-killing bark beetle (Dendroctonus rufipennis Kirby) is phytopathogenic to host trees and may hasten tree decline during colonization by beetles, but defense responses of mature trees to the fungus have not been experimentally examined. To test the hypothesis that interspecific variation in spruce resistance is explained by defense traits we compared constitutive (bark thickness and constitutive resin ducts) and induced defenses (resin flow, monoterpene composition, concentration, phloem lesion formation and traumatic resin ducts) between two sympatric spruces: Engelmann spruce (Picea engelmannii Parry ex Engelm.-a susceptible host) and blue spruce (Picea pungens Engelm.-a resistant host) in response to fungal inoculation. Four central findings emerged: (i) blue spruce has thicker outer bark and thinner phloem than Engelmann spruce, which may restrict fungal access to phloem and result in less beetle-available resource overall; (ii) both spruce species induce monoterpenes in response to inoculation but blue spruce has higher constitutive monoterpene levels, induces monoterpenes more rapidly, and induces higher concentrations over a period of time consistent with spruce beetle attack duration; (iii) Engelmann and blue spruce differed in the monoterpenes they upregulated in response to fungal inoculation: blue spruce upregulated α-pinene, terpinolene and γ-terpinene, but Engelmann spruce upregulated 3-carene and linalool; and (iv) blue spruce has a higher frequency of constitutive resin ducts and produces more traumatic resin ducts in annual growth increments than Engelmann spruce, though Engelmann spruce produces more resin following aseptic wounding or fungal inoculation. These findings suggest that higher constitutive resin duct densities and monoterpene concentrations, as well as the ability to rapidly induce specific monoterpenes in response to L. abietinum inoculation, are phenotypic traits associated with hosts resistant to spruce beetle colonization.

Jasmonate induced terpene-based defense in Pinus massoniana depresses Monochamus alternatus adult feeding.

Symptoms of pine wilt disease (PWD) are caused when pathogenic pine wood nematodes (PWN) invade healthy host trees via wounds created by adult Monochamus alternatus. Exogenous methyl jasmonate (MeJA) can trigger terpene-based induced defense in conifers, which is a core part of the conifer defense system. Herein, we hypothesized that the terpene-based plant defense induced by MeJA could negatively affect the feeding behavior of M. alternatus adults, and may contribute to a new strategy in the control of PWD. The feeding area for M. alternatus adults feeding on MeJA-treated seedlings was significantly smaller compared with control seedlings. However, no significant difference was detected in the enzyme activities in the midgut of beetles that had fed on these seedlings. Terpenoids were mainly accumulated in traumatic resin duct, whereas the constitutive resin duct accumulated only diterpenoids. Correspondingly, large-scaled responses at the transcriptional level mainly focused on terpenoid and phenolic biosynthesis in the defending trees. Breeding tree species with a high resin yield may contribute to control of the spread of PWD by suppressing the feeding of M. alternatus adults. Transcriptome sequencing results provided abundant information for further breeding of highly resistant trees. Based on these findings, a potential push-pull strategy for the control of M. alternatus was discussed.

Tree resin flow dynamics during an experimentally induced attack by Ips avulsus, I. calligraphus, and I. grandicollis

The success of tree colonization by bark beetles depends on their ability to overcome host tree defenses, including resin exudation and toxic chemicals, which deter bark beetle colonization. Resin defenses during insect outbreaks are challenging to study in situ, as outbreaks are stochastic events that progress quickly and thus preclude the establishment of baseline observations of non-infested controls. We use synthetic aggregation pheromones to demonstrate that confined Ips bark beetle herbivory can be successfully initiated to provide opportunities for studying interactions between bark beetles and their hosts, including the dynamics of constitutive and induced resin exudation. In Pinus taeda L. plantations between 12 and 19 years old in North and South Carolina, U.S., trees were affixed with pheromone lures, monitored for evidence of bark beetle attacks, and resin samples were collected throughout the growing season. Baiting increased beetle herbivory to an extent sufficient to produce an induced resin response. Attacked trees exuded about three times more resin at some time than control trees. This supports previous work that demonstrated that information on constitutive resin dynamics alone provides an incomplete view of a host tree’s resistance to bark beetle attack.

Dwarf mistletoe infection in jack pine alters growth-defense relationships.

Trees utilize a combination of chemical and anatomical defenses against a myriad of attacking organisms. However, persistent pathogen infection that alters resource acquisition may impact growth and defense relationships, which could have consequences for tree resistance. We characterized systemic chemical and anatomical changes in jack pine (Pinus banksiana) in response to infection by the parasitic plant dwarf mistletoe (Arceuthobium americanum) and identified how the growth-defense relationship is altered due to infection severity. Our study found that the growth and defense relationship in jack pine was altered due to infection and that chemical defenses in the phloem received a relatively higher priority than radial growth and anatomical defenses. Chemical defenses in the phloem had a non-linear relationship with infection severity with increasing concentrations of monoterpenes in trees with moderate infection and decreasing concentrations at high infection. In contrast, both radial growth and vertical resin duct production decreased with increasing infection severity. While constitutive resin duct counts and many monoterpene compound concentrations were positively correlated, this relationship was not maintained in infected trees. Furthermore, radial growth and basal area increment was positively correlated with resin duct production and monoterpene concentration in non-infected trees but had fewer relationships in severely infected trees. We conclude that while both chemical and anatomical defenses may be used as indicators for potential resistance to biotic stress in pines, changes in resource allocation patterns between these defenses after infection will likely have consequences on tree resistance to subsequent biotic attacks.

Resin monoterpene defenses decline within three widespread species of pine (Pinus) along a 1530‐m elevational gradient

AbstractThe elevational gradient in plant defense (EGPD) hypothesis posits that natural enemy pressures increase alongside temperature across elevational climatic gradients, thereby selecting for enhanced defenses at lower elevations while leaving plants less defended at higher elevations. Phylogenetically constrained tests of this hypothesis are uncommon, with tests focused on defenses of mature trees in natural settings being exceedingly rare. In the absence of this information, predicting the spatiotemporal dynamics of forest pests that preferentially attack mature trees is rendered more difficult. Tree properties such as age, growth rate, and size have all been correlated with levels of tree defenses against insect pests. Thus, we sought to disentangle the interacting influences of these properties from possible elevational climatic effects on monoterpene concentrations, diversity, and intraspecific dissimilarity of constitutive resin within three widespread pines (Pinus contorta, Pinus ponderosa, and Pinus flexilis) across a 1530‐m elevational transect in the Rocky Mountains, Colorado, USA. Overall, we found some support for the EGPD hypothesis within all three of the pine species studied. However, while elevation and tree properties were strong predictors of the variation in monoterpene concentration and moderate predictors of variation in diversity and dissimilarity in P. contorta, they were a relatively weak set of predictors in P. ponderosa and P. flexilis. Significant, (negative) elevational effects were found for total monoterpene concentration in P. contorta and monoterpene diversity in P. ponderosa, while elevation had a significant negative effect on monoterpene concentration and diversity, and a positive effect on dissimilarity in P. flexilis. Nevertheless, tree age, size, and/or growth measures had significant influences on one or more monoterpene measures in all three of the pine species indicating a need to parse effects of these factors within studies of tree defenses across environmental gradients. Our results offer some support for the EGPD hypothesis while suggesting the presence of a variable hierarchy of drivers of pine defense and resistance across congeners in shared environments—a situation that presents challenges for predicting spatiotemporal patterns in secondary chemistry across species from a common set of potential drivers.

Differentiation of persistent anatomical defensive structures is costly and determined by nutrient availability and genetic growth-defence constraints.

Conifers exhibit a number of chemical and anatomical mechanisms to defend against pests and pathogens. Theory predicts an increased investment in plant defences under limited nutrient availability, but while this has been demonstrated for chemical defences, it has rarely been shown for anatomical defensive structures. In a long-lived woody plant, we tested the hypothesis that limited nutrient availability may promote an improved differentiation of persistent anatomical defences. We also hypothesized that the costs of differentiation of those long-term anatomical structures may be determined by genetic constraints on early growth potential. Using Pinus pinaster Ait. juveniles, we performed a greenhouse study with 15 half-sib families subjected to experimental manipulation of phosphorus (P) availability and herbivory-related induced responses. When plants were ∼30 cm high, half of the plant material was treated with methyl jasmonate to induce defences, and 2 weeks later plants were harvested and the abundance of resin canals in the cortex and xylem was assessed. Density of constitutive resin canals in the cortex and the total canal system was ∼1.5-fold higher in plants under limited P availability than in fully fertilized plants. Availability of P did not significantly influence the inducibility of resin canal traits. We found negative genetic correlations between plant growth and the density of constitutive canals in the xylem and total canal system, but only under conditions of limited nutrition. These results demonstrate for the first time that differentiation of constitutive anatomical-based defences is affected by P limitation. Moreover, results also evidence the existence of genetic constraints between plant growth and constitutive defensive investment, where lineages with the highest growth potential showed the lowest investment in constitutive resin canals.

Do water-limiting conditions predispose Norway spruce to bark beetle attack?

Drought is considered to enhance susceptibility of Norway spruce (Picea abies) to infestations by the Eurasian spruce bark beetle (Ips typographus, Coleoptera: Curculionidae), although empirical evidence is scarce. We studied the impact of experimentally induced drought on tree water status and constitutive resin flow, and how physiological stress affects host acceptance and resistance.We established rain-out shelters to induce both severe (two full-cover plots) and moderate (two semi-cover plots) drought stress. In total, 18 sample trees, which were divided equally between the above treatment plots and two control plots, were investigated. Infestation was controlled experimentally using a novel ‘attack box’ method.Treatments influenced the ratios of successful and defended attacks, but predisposition of trees to infestation appeared to be mainly driven by variations in stress status of the individual trees over time. With increasingly negative twig water potentials and decreasing resin exudation, the defence capability of the spruce trees decreased.We provide empirical evidence that water-limiting conditions impair Norway spruce resistance to bark beetle attack. Yet, at the same time our data point to reduced host acceptance byI. typographus with more extreme drought stress, indicated by strongly negative pre-dawn twig water potentials.

Hemlock woolly adelgid (Hemiptera: Adelgidae) induces twig volatiles of eastern hemlock in a forest setting.

The hemlock woolly adelgid (Adelges tsugae Annand) is an invasive species causing high mortality of eastern hemlock (Tsuga canadensis (L) Carriere) in the forests of eastern North America. Recent findings revealed that sapling eastern hemlocks artificially infested with hemlock woolly adelgid in a plantation setting responded to the insect with an array of induced resin volatile changes. Here we determine if eastern hemlocks growing beneath a forest canopy respond to hemlock woolly adelgid infestation with the same patterns of constitutive and inducible volatile resin production as those plantation specimens. We inoculated previously uninfested branches of mature and immature hemlocks in a central New England forest with hemlock woolly adelgid. We then sampled twig tissue of infested and uninfested trees in late spring, early summer, and mid-autumn, after known intervals of adelgid activity when an induced response might be expected. We identified and quantified resin volatiles by gas chromatography-mass spectrometry. Constitutive levels of total monoterpenoids, sesquiterpenoids, and combined resin volatiles were all several-fold more abundant in forest trees than those previously measured in a plantation setting, pointing to further study of the influence of site factors on hemlock volatile production. Hemlock woolly adelgid infestation induced an array of changes in eastern hemlock's volatile profile, including many-fold increases in benzyl alcohol and methyl salicylate accumulation. Despite differences in constitutive concentrations of volatiles between the two sites, our findings verify that hemlock woolly adelgid elicits patterns of resin volatile induction in forest-grown eastern hemlocks quite similar to those previously observed in plantation grown trees.

Discovering candidate genes that regulate resin canal number in Pinus taeda stems by integrating genetic analysis across environments, ages, and populations.

Genetically improving constitutive resin canal development in Pinus stems may enhance the capacity to synthesize terpenes for bark beetle resistance, chemical feedstocks, and biofuels. To discover genes that potentially regulate axial resin canal number (RCN), single nucleotide polymorphisms (SNPs) in 4027 genes were tested for association with RCN in two growth rings and three environments in a complex pedigree of 520 Pinus taeda individuals (CCLONES). The map locations of associated genes were compared with RCN quantitative trait loci (QTLs) in a (P. taeda × Pinus elliottii) × P. elliottii pseudo-backcross of 345 full-sibs (BC1). Resin canal number was heritable (h(2) ˜ 0.12-0.21) and positively genetically correlated with xylem growth (rg ˜ 0.32-0.72) and oleoresin flow (rg ˜ 0.15-0.51). Sixteen well-supported candidate regulators of RCN were discovered in CCLONES, including genes associated across sites and ages, unidirectionally associated with oleoresin flow and xylem growth, and mapped to RCN QTLs in BC1. Breeding is predicted to increase RCN 11% in one generation and could be accelerated with genomic selection at accuracies of 0.45-0.52 across environments. There is significant genetic variation for RCN in loblolly pine, which can be exploited in breeding for elevated terpene content.

Delivering Sitka spruce with resistance against white pine weevil in British Columbia, Canada

The Sitka spruce (Picea sitchensis [Bong.] Carr) breeding program for resistance against the white pine weevil Pissodes strobi Peck (Coleoptera: Curculionidae) is arguably one of the most successful pest resistance breeding programs for plantation forest species in North America, with a substantial proportion of the planting stock in BC and Washington State currently coming from this breeding program. Using conventional selection and breeding, and by screening Sitka spruce populations using artificial weevil infestations, we identified sources of heritable and stable weevil resistance. We also used this program to investigate potential causes behind this resistance and identified several heritable resistance mechanisms, including anatomical characteristics, such as constitutive resin canals and sclereid cells in the bark, terpene defenses and variation in tree phenology. We concluded that resistance is conferred by a suite of traits whose composition varies among resistant sources. In addition, we evaluated the efficiency of screening for resistance using weevil population enhancement as a screening method. Our results culminated in the establishment of seed orchards, and the availability of resistant seed that is contributing to the return of Sitka spruce as a species of choice in coastal British Columbia.

Resistance of Sitka spruce (Picea sitchensis (Bong.) Carr.) to white pine weevil (Pissodes strobi Peck): characterizing the bark defence mechanisms of resistant populations

It has long been known that strong expressions of resistance to the white pine weevil (Pissodes strobi Peck) exist in certain Sitka spruce (Picea sitchensis (Bong.) Carr.) populations, particularly among trees originating from the Fraser Valley and the Qualicum area of British Columbia (BC). In this paper, we characterize how resistance is manifested in these known resistant populations. Specifically, using cloned individuals, we investigated resistant traits associated with repellency, constitutive resin canals (CRC) and sclereid or stone cells. Results indicate significant population differences in the level of these traits between these two populations and susceptible populations. Fraser Valley populations had four times the sclereid density of susceptible populations. Although the Big Qualicum (East Vancouver Island) population had the same high resistance as the Haney (Fraser Valley) population, it was expressed primarily through increased CRC. Sclereid cell density had the strongest correlation to weevil attack followed by CRC. We discuss pathways by which two distinct resistant populations may have developed in this high weevil hazard region of south-west BC.

Differences in the constitutive terpene profile of lodgepole pine across a geographical range in British Columbia, and correlation with historical attack by mountain pine beetle

AbstractThe mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), is a destructive insect pest in western Nearctic conifer forests. Currently, British Columbia, Canada, is experiencing the largest recorded outbreak of this insect, including areas that historically have had low climatic suitability for it. We analyzed 26 constitutive resin terpenes in phloem samples from British Columbia lodgepole pine (Pinus contorta) populations to test for differential resistance to mountain pine beetle attack, based upon the likelihood of previous exposure to mountain pine beetle. We assessed sampled trees for number of mountain pine beetle attacks, number of pupal chambers, and tree survival the following spring. Significant differences were found when levels of certain terpenes in lodgepole pine populations that had likely experienced substantial mountain pine beetle infestations in the past were compared with those in populations that likely had not experienced large outbreaks of mountain pine beetle. Although we expected southern pine populations to contain more total terpenes than northern populations, owing to higher historical exposure to the beetle, the converse was found. Northern populations generally had higher levels of constitutive terpenes and beetle attack than southern populations. Because several terpenes are kairomones to the mountain pine beetle and also serve as precursors for the synthesis of pheromones, the lower levels of terpenes expressed by lodgepole pines from the historical range of the mountain pine beetle may render them less chemically perceptible to foraging beetles.

Resin flow responses to fertilization, wounding and fungal inoculation in loblolly pine (Pinus taeda) in North Carolina

Resin flow is the primary means of natural defense against southern pine beetle (Dendroctonus frontalis Zimm.), the most important insect pest of Pinus spp. in the southern United States. As a result, factors affecting resin flow are of interest to researchers and forest managers. We examined the influence of fertilization, artificial wounding and fungal inoculation on resin flow in 6- and 12-year-old stands of loblolly pine (Pinus taeda L.) and determined the extent of that influence within and above the wounded stem area and through time. Fertilization increased constitutive resin flow, but only the younger trees sustained increased resin flow after wounding and inoculation treatments. An induced resin flow response occurred between 1 and 30 days after wounding and inoculation treatments. Wounding with inoculation resulted in greater resin flow than wounding alone, but increasing amounts of inoculum did not increase resin flow. Increased resin flow (relative to controls) lasted for at least 90 days after wounding and inoculation. This increase appeared to be limited to the area of treatment, at least in younger trees. The long-lasting effects of fungal inoculation on resin flow, as well as the response to fertilization, suggest that acquired resistance through induced resin flow aids in decreasing susceptibility of loblolly pine to southern pine beetle.

Why does longleaf pine have low susceptibility to southern pine beetle?

Pine forests throughout the world are subject to disturbance from tree-killing bark beetles, but pine species differ in their susceptibilities. In the southeastern United States, Pinus palustris Mill. suffers far less mortality from the southern pine beetle, Dendroctonus frontalis Zimmermann, than do its sympatric congeners. We tested the commonly invoked hypothesis that P. palustris has relatively low susceptibility because it has higher oleoresin flow than other pines, especially Pinus taeda L. However, seven studies in three states over 6 years refuted the hypothesis that P. palustris and P. taeda differ in their constitutive resin flow or in their capacity to replace resin depleted by either experimental wounding or natural beetle attacks. Additionally, surveys of natural beetle attacks revealed that P. taeda and P. palustris were equally likely to be attacked and killed when they cooccurred in front of growing infestations. Thus, the relative susceptibility of these two species changes with the spatial scale at which they are mixed, and the strong landscape-scale pattern of low mortality in P. palustris is not because individual trees are physiologically less susceptible. Ultimately, the conspicuous differential impact of D. frontalis on P. taeda and P. palustris may be the product of coevolution between tree defenses and beetle behavior.

Effects of mass inoculation on induced oleoresin response in intensively managed loblolly pine

Oleoresin flow is an important factor in the resistance of pines to attack by southern pine beetle, Dendroctonus frontalis Zimm., and its associated fungi. Abiotic factors, such as nutrient supply and water relations, have the potential to modify this plant-insect-fungus interaction; however, little is known of the effects of inoculation with beetle-associated fungi on oleoresin flow. We observed that constitutive and induced resin yield in loblolly pine, Pinus taeda L., were affected by either fungal inoculation (with the southern pine beetle-associated fungus Ophiostoma minus (Hedgcock) H. & P. Sydow) or silvicultural treatment. The effects of mass wounding (400 wounds m(-2)) and mass wounding and inoculation with O. minus were assessed by comparison with untreated (control) trees. The treatments were applied to trees in a 2 x 2 factorial combination of fertilizer and irrigation treatments. Fertilization did not significantly affect constitutive resin yield. Even as long as 105 days post-treatment, however, mass-inoculated trees produced higher induced resin yields than control or wounded-only trees, indicating a localized induced response to fungal inoculation. We noted no systemic induction of host defenses against fungal colonization. Although beetles attacking previously attacked trees face a greater resinous response from their host than beetles attacking trees that had not been previously attacked, the effect of an earlier attack may not last more than one flight season. Despite mass inoculations, O. minus did not kill the host trees, suggesting that this fungus is not a virulent plant pathogen.