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Concept: Pinus classification


Mongolian Scots pine (Pinus sylvestris var. mongolica) is one of the principal tree species in the network of Three-North Shelterbelt for windbreak and sand stabilisation in China. The functions of shelterbelts are highly correlated with the architecture and eco-physiological processes of individual tree. Thus, model-assisted analysis of canopy architecture and function dynamic in Mongolian Scots pine is of value for better understanding its role and behaviour within shelterbelt ecosystems in these arid and semiarid regions. We present here a single-tree functional and structural model, derived from the GreenLab model, which is adapted for young Mongolian Scots pines by incorporation of plant biomass production, allocation, allometric rules and soil water dynamics. The model is calibrated and validated based on experimental measurements taken on Mongolian Scots pines in 2007 and 2006 under local meteorological conditions. Measurements include plant biomass, topology and geometry, as well as soil attributes and standard meteorological data. After calibration, the model allows reconstruction of three-dimensional (3D) canopy architecture and biomass dynamics for trees from one- to six-year-old at the same site using meteorological data for the six years from 2001 to 2006. Sensitivity analysis indicates that rainfall variation has more influence on biomass increment than on architecture, and the internode and needle compartments and the aboveground biomass respond linearly to increases in precipitation. Sensitivity analysis also shows that the balance between internode and needle growth varies only slightly within the range of precipitations considered here. The model is expected to be used to investigate the growth of Mongolian Scots pines in other regions with different soils and climates.

Concepts: Precipitation, Soil, Tree, Scots Pine, Pine, Pinus classification, Biomass, Pinus


Diterpene resin acids (DRAs) are major components of pine (Pinus spp.) oleoresin. They play critical roles in conifer defense against insects and pathogens and as a renewable resource for industrial bioproducts. The core structures of DRAs are formed in secondary (i.e. specialized) metabolism via cycloisomerization of geranylgeranyl diphosphate (GGPP) by diterpene synthases (diTPSs). Previously described gymnosperm diTPSs of DRA biosynthesis are bifunctional enzymes that catalyze the initial bicyclization of GGPP followed by rearrangement of a (+)-copalyl diphosphate intermediate at two discrete class II and class I active sites. In contrast, similar diterpenes of gibberellin primary (i.e. general) metabolism are produced by the consecutive activity of two monofunctional class II and class I diTPSs. Using high-throughput transcriptome sequencing, we discovered 11 diTPS from jack pine (Pinus banksiana) and lodgepole pine (Pinus contorta). Three of these were orthologous to known conifer bifunctional levopimaradiene/abietadiene synthases. Surprisingly, two sets of orthologous PbdiTPSs and PcdiTPSs were monofunctional class I enzymes that lacked functional class II active sites and converted (+)-copalyl diphosphate, but not GGPP, into isopimaradiene and pimaradiene as major products. Diterpene profiles and transcriptome sequences of lodgepole pine and jack pine are consistent with roles for these diTPSs in DRA biosynthesis. The monofunctional class I diTPSs of DRA biosynthesis form a new clade within the gymnosperm-specific TPS-d3 subfamily that evolved from bifunctional diTPS rather than monofunctional enzymes (TPS-c and TPS-e) of gibberellin metabolism. Homology modeling suggested alterations in the class I active site that may have contributed to their functional specialization relative to other conifer diTPSs.

Concepts: Pinophyta, Pine, Pinus classification, Resin, Lodgepole Pine, Conifer cone, Pinus, Abietic acid


Fossil records indicate that the genus Pinus L. split into two subgenera by the Late Cretaceous, although subgenus Strobus (D. Don) Lemmon is less well documented than subgenus Pinus L., especially in eastern Asia. In this paper, Pinus maomingensis sp. nov. is established based on a compressed seed cone from the upper Eocene of the Maoming Basin of southern China. This species is attributed to genus Pinus, subgenus Strobus, section Quinquefoliae Duhamel, subsection Strobus Loudon based on the combination of morphological characters obtained from the cone scales, specifically from the terminal umbo, rhombic apophysis, and cuticle structure. Associated fascicles of needle leaves with deciduous sheaths and bulbous bases are recognized as Pinus sp. and also represent Pinus subgenus Strobus. This new discovery from the Maoming Basin constitutes the first megafossil record of subgenus Strobus from southern China and implies that the members of this subgenus arrived in the southern region of China by the late Eocene. The extant species of subgenus Strobus are mainly distributed in northern temperate and tropical to subtropical mountainous regions. We propose that the Maoming Basin was adjacent to a mountainous region during the late Eocene.

Concepts: Mammal, Leaf, Pine, Pinus classification, Eocene, Conifer cone, Pinus


In response to climate warming, subalpine treelines are expected to move up in elevation since treelines are generally controlled by growing season temperature. Where treeline is advancing, dispersal differences and early life stage environmental tolerances are likely to affect how species expand their ranges. Species with an establishment advantage will colonize newly available habitat first, potentially excluding species that have slower establishment rates. Using a network of plots across five mountain ranges, we described patterns of upslope elevational range shift for the two dominant Great Basin sub-alpine species, limber pine and Great Basin bristlecone pine. We found that the Great Basin treeline for these species is expanding upslope with a mean vertical elevation shift of 19.1 m since 1950, which is lower than what we might expect based on temperature increases alone. The largest advances were on limber pine-dominated granitic soils, on west aspects, and at lower latitudes. Bristlecone pine juveniles establishing above treeline share some environmental associations with bristlecone adults. Limber pine above-treeline juveniles, in contrast, are prevalent across environmental conditions and share few environmental associations with limber pine adults. Strikingly, limber pine is establishing above treeline throughout the region without regard to site characteristic such as soil type, slope, aspect, or soil texture. Though limber pine is often rare at treeline where it coexists with bristlecone pine, limber pine juveniles dominate above treeline even on calcareous soils that are core bristlecone pine habitat. Limber pine is successfully “leap-frogging” over bristlecone pine, probably because of its strong dispersal advantage and broader tolerances for establishment. This early-stage dominance indicates the potential for the species composition of treeline to change in response to climate change. More broadly, it shows how species differences in dispersal and establishment may result in future communities with very different specific composition. This article is protected by copyright. All rights reserved.

Concepts: Climate, Soil, Climate change, Pinus classification, Pinus, Whitebark Pine, Tree line, Bristlecone pine


Warming climate has increased access of native bark beetles to high-elevation pines that historically received only intermittent exposure to these tree-killing herbivores. Here we show that a dominant, relatively naïve, high-elevation species, whitebark pine, has inferior defenses against mountain pine beetle compared with its historical lower-elevation host, lodgepole pine. Lodgepole pines respond by exuding more resin and accumulating higher concentrations of toxic monoterpenes than whitebark pine, where they co-occur. Furthermore, the chemical composition of whitebark pine appears less able to inhibit the pheromonal communication beetles use to jointly overcome tree defenses. Despite whitebark pine’s inferior defenses, beetles were more likely to attack their historical host in mixed stands. This finding suggests there has been insufficient sustained contact for beetles to alter their complex behavioral mechanisms driving host preference. In no-choice assays, however, beetles readily entered and tunneled in both hosts equally, and in stands containing less lodgepole pine, attacks on whitebark pines increased. High-elevation trees in pure stands may thus be particularly vulnerable to temperature-driven range expansions. Predators and competitors were more attracted to volatiles from herbivores attacking their historical host, further increasing risk in less coevolved systems. Our results suggest cold temperatures provided a sufficient barrier against herbivores for high-elevation trees to allocate resources to other physiological processes besides defense. Changing climate may reduce the viability of that evolutionary strategy, and the life histories of high-elevation trees seem unlikely to foster rapid counter adaptation. Consequences extend from reduced food supplies for endangered grizzly bears to altered landscape and hydrological processes.

Concepts: Beetle, Attack, Pine, Pinus classification, Curculionidae, Lodgepole Pine, Pinus, Bark beetle


Until very recently, complete characterization of the megagenomes of conifers has remained elusive. The diploid genome of sugar pine (Pinus lambertiana Dougl.) has a highly repetitive 31 billion base pair genome. With nearly 50% more DNA than the current record holder, Pinus taeda, sugar pine is the largest genome sequenced to date. This genome is the first to be sequenced from the subgenus Strobus, or white pines, a group that is notable for having the largest genomes among the pines. The genome represents a unique opportunity to investigate genome “obesity” in conifers and white pines. The crux of our sequencing approach has been to leverage aspects of conifer biology to reduce the complexity of the assembly problem. Comparative analysis of P. lambertiana and P. taeda reveals new insights on the conservation, age, and diversity of the highly abundant transposable elements, the primary factor determining genome size. Like most North American white pines the principal pathogen of P. lambertiana is white pine blister rust (Cronartium ribicola J.C. Fischer ex Raben.). Identification of the candidate genes for resistance to this pathogen is great ecological importance. The genome afforded us the opportunity to make substantial progress on locating the major dominant gene for simple resistance hypersensitive response, Cr1 We describe new markers and gene annotation that are both tightly linked to Cr1 in a mapping population and associated to Cr1 in unrelated individuals sampled throughout the range, creating a solid foundation for future mapping. This genomic variation and annotated candidate genes characterized in our study of Cr1 region are resources for future marker-assisted breeding efforts as well as investigations of fundamental mechanisms of invasive disease and evolutionary response.

Concepts: DNA, Gene, Genetics, Evolution, Genome, Pine, Pinus classification, Western White Pine


Extracts from pine tree bark containing a variety of flavonoids have been used in traditional medicine. Pycnogenol is a proprietary bark extract of the French maritime pine tree (Pinus pinaster ssp. atlantica) that exerts antioxidative, anti-inflammatory, and anti-platelet effects. However, the effects of Pycnogenol on endothelial dysfunction, a precursor of atherosclerosis and cardiovascular events, remain still elusive.

Concepts: Atherosclerosis, Coronary artery disease, Heart, Blood vessel, Pine, Pinus classification, Pinus, BARK


The toxicity of red pine needle hydrodistillate (RPN-HD), 19 RPN-HD constituents and another 12 structurally related compounds and the control efficacy of four experimental spray formulations containing RPN-HD (0.5. 1, 2 and 3% sprays) to adult Dermatophagoides farinae were evaluated.

Concepts: House dust mite, Pine, Pinus classification, Pinus


To develop safe and effective methods to protect whitebark pines, Pinus albicaulis Engelmann, and limber pines, Pinus flexilis James, from attack by mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), we compared verbenone and verbenone plus green leaf volatiles (GLVs) for prevention of beetle attack. We used two strategies: area-wide protection where semiochemical-releasing flakes are dispersed over the forest floor, and individual tree tests where flakes are applied to tree trunks. The area-wide bioassays were conducted by applying verbenone- and GLV-releasing flakes without stickers to the forest floor on 0.81-ha plots dominated by whitebark pines in the State of Washington with four replicates. We conducted individual tree bioassays by applying the same formulations with stickers to whitebark and limber pines in Montana and Colorado, respectively. In all three situations, both verbenone-alone and verbenone plus GLVs significantly increased the proportion of trees escaping mass attack by beetles, but the two formulations were not significantly different from one another. Despite a lack of significance at a Bonferroni-adjusted α = 0.05, adding GLVs gave slightly greater absolute levels of tree protection in most cases. Monitoring traps placed in the area-wide treatments in Washington showed similar outcomes for numbers of beetles trapped: both treatments had significantly fewer beetles than controls, and they were not significantly different from one another. At peak flight, however, plots with GLVs combined with verbenone had roughly 40% fewer beetles than plots with verbenone alone. GLVs are considerably cheaper than verbenone, so tests of higher application rates may be warranted to achieve enhanced tree protection at reasonable cost.

Concepts: Beetle, Pine, Pinus classification, Curculionidae, Pinus, Mountain pine beetle, Bark beetle, Tree line


Plants have an accumulative response to heavy metals present in soils or deposited from airborne sources of emissions. Therefore, their tissues are very often used in studies of heavy metal contamination originating from different sources as a bioindicator of environmental pollution. This research was undertaken to examine accumulation capacities of Pb, Zn, Cd, Cu and Cr in washed and unwashed needles of Scots pine (Pinus sylvestris L.) and leaves of silver birch (Betula pendula Roth) growing in a contaminated area. We collected needles of Scots pine and leaves of silver birch in an area around a sedimentation pond and metallurgic plant processing Pb and Zn ores near Olkusz, Poland. Concentrations of heavy metals, which have been linked with exposure to emissions, were determined from foliar samples collected at 33 sites. These sites were established at various distances (0.5-3.6 km) from the pond and metallurgic plant so as to identify the predominant accumulative response of plants. Spatial gradients for Pb and Zn were calculated using an ordinary kriging interpolation algorithm. A spatial pattern was identified by a GIS method to visualize maps over the Pb-Zn ore mining area. The accumulation of Zn (R(2) = 0.74, p < 0.05) and Pb (R(2) = 0.85, p < 0.01) in plant tissues correlated with soil concentrations. This tendency was not found in the case of Cu, Cd and Cr.

Concepts: Zinc, Lead, Birch, Heavy metal music, Scots Pine, Pine, Pinus classification, Betula pendula