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Concept: Conifer cone


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


Asymmetry in motor patterns is present in a wide variety of animals. Many lateralized behaviors seem to depend on brain asymmetry, as it is the case of different tasks associated to food handling by several bird and mammal species. Here, we analyzed asymmetry in handling behavior of pine cones by red squirrels (Sciurus vulgaris). Red squirrels devote most of their daily activity to feeding, thus this species constitutes an appropriate model for studying asymmetry in food processing. We aimed to explore 1) the potential lateralization in handling of pine cones by squirrels, 2) the dominant pattern for this behavior (left- vs. right-handed), and 3) whether this pattern varies among populations and depending on the pine tree species available. Results revealed that red squirrels handle pine cones in an asymmetrical way, and that direction of asymmetry varies among populations and seems to be determined more by local influences rather than by the pine tree species.

Concepts: Mammal, Pinophyta, Standard Model, Pine, Conifer cone, Lateralization of brain function, Eastern Gray Squirrel, Tree squirrel


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



When confronted with an adaptive challenge, such as extreme temperature, closely related species frequently evolve similar phenotypes using the same genes. Although such repeated evolution is thought to be less likely in highly polygenic traits and distantly related species, this has not been tested at the genome scale. We performed a population genomic study of convergent local adaptation among two distantly related species, lodgepole pine and interior spruce. We identified a suite of 47 genes, enriched for duplicated genes, with variants associated with spatial variation in temperature or cold hardiness in both species, providing evidence of convergent local adaptation despite 140 million years of separate evolution. These results show that adaptation to climate can be genetically constrained, with certain key genes playing nonredundant roles.

Concepts: DNA, Gene, Genetics, Evolution, Biology, Organism, Pine, Conifer cone


We investigate the utility and scalability of new read cloud technologies to improve the draft genome assemblies of the colossal and largely repetitive genomes of conifers. Synthetic long read technologies existed in various forms as a means of reducing complexity and resolving repeats since the outset of genome assembly. Recently, technologies that combine sub-haploid pools of high molecular weight DNA with barcoding on a massive scale have brought new efficiencies to sample preparation and data generation. When combined with inexpensive light shotgun sequencing the resulting data can be used to scaffold large genomes. The protocol is efficient enough to consider routinely for even the largest genomes. Conifers represent the largest reference genome projects executed to date. The largest of these, is that of the conifer Pinus lambertiana (sugar pine), with a genome size of 31 billion bp. In this paper, we report on the molecular and computational protocols for scaffolding the P. lambertiana genome using the library technology from 10X Genomics. At 247 thousand base pairs, the NG50 of the existing reference sequence is the highest scaffold contiguity among the currently published conifer assemblies; this new assembly’s NG50 is 1.94 million base pairs, an eightfold increase.

Concepts: DNA, Human genome, Human Genome Project, Genome, Pinophyta, Pine, Conifer cone, Pinaceae


Widespread tree mortality caused by outbreaks of native bark beetles (Circulionidae: Scolytinae) in recent decades has raised concern among scientists and forest managers about whether beetle outbreaks fuel more ecologically severe forest fires and impair postfire resilience. To investigate this question, we collected extensive field data following multiple fires that burned subalpine forests in 2011 throughout the Northern Rocky Mountains across a spectrum of prefire beetle outbreak severity, primarily from mountain pine beetle (Dendroctonus ponderosae). We found that recent (2001-2010) beetle outbreak severity was unrelated to most field measures of subsequent fire severity, which was instead driven primarily by extreme burning conditions (weather) and topography. In the red stage (0-2 y following beetle outbreak), fire severity was largely unaffected by prefire outbreak severity with few effects detected only under extreme burning conditions. In the gray stage (3-10 y following beetle outbreak), fire severity was largely unaffected by prefire outbreak severity under moderate conditions, but several measures related to surface fire severity increased with outbreak severity under extreme conditions. Initial postfire tree regeneration of the primary beetle host tree [lodgepole pine (Pinus contorta var. latifolia)] was not directly affected by prefire outbreak severity but was instead driven by the presence of a canopy seedbank and by fire severity. Recent beetle outbreaks in subalpine forests affected few measures of wildfire severity and did not hinder the ability of lodgepole pine forests to regenerate after fire, suggesting that resilience in subalpine forests is not necessarily impaired by recent mountain pine beetle outbreaks.

Concepts: Beetle, Pine, Pinus classification, Lodgepole Pine, Conifer cone, Pinus, Mountain pine beetle, Bark beetle


Escalating wildfire in subalpine forests with stand-replacing fire regimes is increasing the extent of early-seral forests throughout the western USA. Post-fire succession generates the fuel for future fires, but little is known about fuel loads and their variability in young post-fire stands. We sampled fuel profiles in 24-year-old post-fire lodgepole pine (Pinus contorta var. latifolia) stands (n = 82) that regenerated from the 1988 Yellowstone Fires to answer three questions. (1) How do canopy and surface fuel loads vary within and among young lodgepole pine stands? (2) How do canopy and surface fuels vary with pre- and post-fire lodgepole pine stand structure and environmental conditions? (3) How have surface fuels changed between eight and 24 years post-fire? Fuel complexes varied tremendously across the landscape despite having regenerated from the same fires. Available canopy fuel loads and canopy bulk density averaged 8.5 Mg/ha (range 0.0-46.6) and 0.24 kg/m(3) (range: 0.0-2.3), respectively, meeting or exceeding levels in mature lodgepole pine forests. Total surface-fuel loads averaged 123 Mg/ha (range: 43-207), and 88% was in the 1,000-h fuel class. Litter, 1-h, and 10-h surface fuel loads were lower than reported for mature lodgepole pine forests, and 1,000-h fuel loads were similar or greater. Among-plot variation was greater in canopy fuels than surface fuels, and within-plot variation was greater than among-plot variation for nearly all fuels. Post-fire lodgepole pine density was the strongest positive predictor of canopy and fine surface fuel loads. Pre-fire successional stage was the best predictor of 100-h and 1,000-h fuel loads in the post-fire stands and strongly influenced the size and proportion of sound logs (greater when late successional stands had burned) and rotten logs (greater when early successional stands had burned). Our data suggest that 76% of the young post-fire lodgepole pine forests have 1,000-h fuel loads that exceed levels associated with high-severity surface fire potential, and 63% exceed levels associated with active crown fire potential. Fire rotations in Yellowstone National Park are predicted to shorten to a few decades and this prediction cannot be ruled out by a lack of fuels to carry repeated fires.

Concepts: Pine, Pinus classification, Yellowstone National Park, Lodgepole Pine, Conifer cone, Pinus, Wildfire, Blue stain fungus


In the absence of broad-scale disturbance, many temperate coniferous forests experience successful seedling establishment only when abundant seed production coincides with favorable climate. Identifying the frequency of past establishment events and the climate conditions favorable for seedling establishment is essential to understanding how climate warming could affect the frequency of future tree establishment events and therefore future forest composition or even persistence of a forest cover. In the southern Rocky Mountains, USA, research on the sensitivity of establishment of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa)-two widely distributed, co-occurring conifers in North America-to climate variability has focused on the alpine treeline ecotone, leaving uncertainty about the sensitivity of these species across much of their elevation distribution. We compared annual germination dates for >450 Engelmann spruce and >500 subalpine fir seedlings collected across a complex topographic-moisture gradient to climate variability in the Colorado Front Range. We found that Engelmann spruce and subalpine fir established episodically with strong synchrony in establishment events across the study area. Broad-scale establishment events occurred in years of high soil moisture availability, which were characterized by above-average snowpack and/or cool and wet summer climatic conditions. In the recent half of the study period (1975-2010), a decrease in the number of fir and spruce establishment events across their distribution coincided with declining snowpack and a multi-decadal trend of rising summer temperature and increasing moisture deficits. Counter to expected and observed increases in tree establishment with climate warming in maritime subalpine forests, our results show that recruitment declines will likely occur across the core of moisture-limited subalpine tree ranges as warming drives increased moisture deficits.

Concepts: Climate, Climate change, Seed, Spruce, Pinophyta, Fir, Conifer cone, Pinaceae


Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long-term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought-tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed-species stands along an altitudinal gradient (400-1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought-prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change. This article is protected by copyright. All rights reserved.

Concepts: Europe, Douglas-fir, Spruce, Pinophyta, Fir, Christmas tree, Conifer cone, Pinaceae