Search

According to the forest resources inventory data for different periods and the latest estimation parameters of forest carbon reserves in China, the carbon reserves and carbon density of forest biomass in the Tibet Autonomous Region from 1999 to 2019 were estimated using the IPCC international carbon reserves estimation model. The results showed that, during the past 20 years, the forest area, forest stock, and biomass carbon storage in Tibet have been steadily increasing, with an average annual increase of 1.85×104 hm2, 0.033×107 m3, and 0.22×107 t, respectively. Influenced by geographical conditions and the natural environment, the forest area and biomass carbon storage gradually increased from the northwest to the southeast, particularly in Linzhi and Changdu, where there are many primitive forests, which serve as important carbon sinks in Tibet. In terms of the composition of tree species, coniferous forests are dominant in Tibet, particularly those containing Abies fabri, Picea asperata, and Pinus densata, which comprise approximately 45% of the total forest area in Tibet. The ecological location of Tibet has resulted in the area being dominated by shelter forest, comprising 68.76% of the total area, 64.72% of the total forest stock, and 66.34% of the total biomass carbon reserves. The biomass carbon storage was observed to first increase and then decrease with increasing forest age, which is primarily caused by tree growth characteristics. In over-mature forests, trees' photosynthesis decreases along with their accumulation of organic matter, and the trees can die. In addition, this study also observed that the proportion of mature and over-mature forest in Tibet is excessively large, which is not conducive to the sustainable development of forestry in the region. This problem should be addressed in future management and utilization activities., Competing Interests: The authors have declared that no competing interests exist.

Inbreeding depression (ID) is a fundamental selective pressure that shapes mating systems and population genetic structures in plants. Although it has been shown that ID varies over the life stages of shorter-lived plants, less is known about how the fitness effects of inbreeding vary across life stages in long-lived species. We conducted a literature survey in the Pinaceae, a tree family known to harbour some of the highest mutational loads ever reported. Using a meta-regression model, we investigated distributions of inbreeding depression over life stages, adjusting for effects of inbreeding levels and the genetic differentiation of populations within species. The final dataset contained 147 estimates of ID across life stages from 41 studies. 44 Fst estimates were collected from 40 peer-reviewed studies for the 18 species to aid genetic differentiation modelling. Partitioning species into fragmented and well-connected groups using Fst resulted in the best way (i.e. trade-off between high goodness-of-fit of the model to the data and reduced model complexity) to incorporate genetic connectivity in the meta-regression analysis. Inclusion of a life stage term and its interaction with the inbreeding coefficient (F) dramatically increased model precision. We observed that the correlation between ID and F was significant at the earliest life stage. Although partitioning of species populations into fragmented and well-connected groups explained little of the between-study heterogeneity, the inclusion of an interaction between life stage and population differentiation revealed that populations with fragmented distributions suffered lower inbreeding depression at early embryonic stages than species with well-connected populations. There was no evidence for increased ID in late life stages in well-connected populations, although ID tended to increase across life stages in the fragmented group. These findings suggest that life stage data should be included in inbreeding depression studies and that inbreeding needs to be managed over life stages in commercial populations of long-lived plants.

Consequences of climate change will severely affect forest ecosystems in the near future, yet our understanding of how and why trees are responding to their abiotic environment is still limited. Intra-specific variation (ITV) in the growth response of trees to warming and drought has been widely neglected so far, but could play a key role for adapting forests to future climate conditions. We analyzed tree rings from four conifers (Picea abies, Abies alba, Larix decidua, Pseudotsuga menziesii) regarding their intra-specific adaptation potential when trees are growing at the warm and dry margins of species distributions. Our study comprises data from four common garden experiments (45 provenances and a total of 743 trees) and assessed growth response at different temporal scales from decades (long-term) to only a few event years (short-term) and finally for density fluctuations within one year (sudden response). We observed significant variation among provenances at all time-scales, but with varying degree among species. However, variation in short-term response (drought years) was remarkably unstable across all species, when the seasonal variation of drought occurrence was considered. Silver-fir and Douglas-fir showed significant associations between seed-source climate and growth response as well as trade-offs between early- and latewood growth reaction which strongly suggests that growth responses are adaptive. Intra-specific variation in conifers in response to drought will probably be sufficient to mitigate climate change consequences on forest growth, but growth-environment interactions as well as dependencies between temporal scales could create major pitfalls for adaptive forest management in the future., (Copyright © 2019 The Authors. Published by Elsevier B.V. All rights reserved.)

Seed dispersal is critical for plants, but the evolution of mechanisms that actually release seeds from their parents is not well understood. We use the reproductive cones of conifers, specifically the Pinaceae clade, to explore the factors driving the evolution of different release mechanisms in plants. We combine comparative anatomical and phylogenetic analyses to test whether fundamental trade-offs in the mechanical and hydraulic properties of vasculature underlie the evolution of two seed release mechanisms: cone scale flexion and cone scale shedding. We then test whether these mechanisms are linked with differences in seed size, dispersal syndrome and reproductive allocation. Cone scale xylem in flexing species is tough, but poorly conductive. Xylem in shedding species is less extensive, fragile and highly conductive; its thin-walled tracheids allow scales to easily fracture at maturity. Shedding is also consistently associated with large, densely packed seeds. Pinaceae cones exploit a well-known trade-off in xylem mechanical strength vs hydraulic efficiency to generate release mechanisms that allow seeds of various sizes to leave the protecting cone. The linkage among release mechanisms, vascular anatomy and seed traits illustrates how a wide variety of selective pressures may influence the function and physiology of reproductive structures., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

The branch xylem of six important Inner Asian southern boreal forest trees was studied for wood-anatomical and hydraulic traits in order to infer the species' drought tolerance from embolism resistance, potential hydraulic conductivity, mean conduit diameters and conduit density. The only studied angiosperm tree, Betula pendula Roth, was much more sensitive to cavitation than all five conifers (evergreen or summer-green), even when using 88% loss of conductivity (P88) in birch, but 50% (P50) in the conifers as critical thresholds. This suggests that pioneer birch forests, which have widely replaced the conifer climax forests after anthropogenic disturbance (e.g., logging, man-made fire), are more vulnerable to climate warming-induced drought than the original conifer forests. In contrast to expectation, the generally more drought-exposed light taiga species (Larix sibirica Ledeb., Pinus sylvestris L.) did not have consistently lower P50 and P88 values than the dark taiga conifers, suggesting that other drought survival traits are equally important. Among the dark-taiga species, only Pinus sibirica Du Tour, but not Abies sibirica Ledeb. and Picea obovata Ledeb., had relatively high P50 values indicating higher vulnerability. In the light-taiga forest, P. sylvestris revealed lower embolism resistance than L. sibirica. In the face of rapid climate warming and drying in Inner Asia, the drought survival strategies of southern boreal tree species deserve further intensive study, which should include other drought survival traits., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Resins are plant exudates of economic importance used by plants as defence. They flow out of resin ducts, open and long tube-like intercellular spaces lined by a layer of specialized parenchyma cells, called the epithelium, which secrete resin into the duct lumen. A model that describes resin flow in conifers is presented to investigate how duct structure, resin loading, crystallisation, and viscosity affect flow and could explain differences between species. Considering resin viscosity, the structure of resin ducts, and a pressure-driven resin loading through the duct wall, the unsteady Stokes equation was applied. There is an increase in flow towards the open end that is favoured by the duct geometry. Both flow and pressure depend on the loading mechanism and on the duct resistance, which depends on the duct geometry, viscosity and duct wall permeability to resin. These results confirm previous measurements and observations made on Pinaceae and seem to be physiologically advantageous for the defence role of resin. Understanding of how these physiological and morphological parameters affect resin flow might be useful for selecting varieties and species having a high resin yielding capacity. The model presented in this paper is also applicable to other external secretory systems in plants., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Temperatures are expected to increase over the next century in all terrestrial biomes and particularly in boreal forests, where drought-induced mortality has been predicted to rise. Genomics research is helping to develop hypotheses regarding the molecular basis of drought tolerance and recent work proposed that the osmo-protecting dehydrin proteins have undergone a clade-specific expansion in the Pinaceae, a major group of conifer trees. The objectives of this study were to identify all of the putative members of the gene family, trace their evolutionary origin, examine their structural diversity and test for drought-responsive expression. We identified 41 complete dehydrin coding sequences in Picea glauca, which is four times more than most angiosperms studied to date, and more than in pines. Phylogenetic reconstructions indicated that the family has undergone an expansion in conifers, with parallel evolution implicating the sporadic resurgence of certain amino acid sequence motifs, and a major duplication giving rise to a clade specific to the Pinaceae. A variety of plant dehydrin structures were identified with variable numbers of the A-, E-, S- and K-segments and an N-terminal (N1) amino acid motif including assemblages specific to conifers. The expression of several of the spruce dehydrins was tissue preferential under non-stressful conditions or responded to water stress after 7-18 days without watering, reflecting changes in osmotic potential. We found that dehydrins with N1 K2 and N1 AESK2 sequences were the most responsive to the lack of water. Together, the family expansion, drought-responsive expression and structural diversification involving loss and gain of amino acid motifs suggests that subfunctionalization has driven the diversification seen among dehydrin gene duplicates. Our findings clearly indicate that dehydrins represent a large family of candidate genes for drought tolerance in spruces and in other Pinaceae that may underpin adaptability in spatially and temporally variable environments.

Mast seeding, or masting, is the highly variable and spatially synchronous production of seeds by a population of plants. The production of variable seed crops is typically correlated with weather, so it is of considerable interest whether global climate change has altered the variability of masting or the size of masting events. We compiled 1086 datasets of plant seed production spanning 1900-2014 and from around the world, and then analysed whether the coefficient of variation (CV) in seed set, a measure of masting, increased over time. Over this 115-year period, seed set became more variable for plants as a whole and for the particularly well-studied taxa of conifers and oaks. The increase in CV corresponded with a decrease in the long-term mean of seed set of plant species. Seed set CV increased to a greater degree in plant taxa with a tendency towards masting. Seed set is becoming more variable among years, especially for plant taxa whose masting events are known to affect animal populations. Such subtle change in reproduction can have wide-ranging effects on ecosystems because seed crops provide critical resources for a wide range of taxa and have cascading effects throughout food webs., (© 2017 The Author(s).)

Spruce can grow at an extra low temperature (LT), and is inferred with important antifreezing gene resources. The research here identified 4 different spruce varieties, named as PicW1, PicW2, PicM and PicK. Sequence alignment showed base-substitution and deficiency mutations among them with sequence identity between 97.61% and 99.25%. Each gene was transferred into E. coli, where protein was induced by IPTG (isopropyl-β-d-thiogalactoside). Strains cultured at -5°C showed the lethal dose 50% (LD-50) between 53h and 57h for the transgenic strains, but 35h for the control. Strains cultivated at -20°C showed the LD-50 between 38h and 44h for the transgenic strains, but 25h for the control. Further, the soluble gene proteins were extracted and purified for Differential Scanning Calorimeter (DSC) test, which showed characteristic thermal hysteresis (TH) value of 0.77°C (PicW1), 0.78°C (PicW2), 0.72°C (PicM), and 0.86°C (PicK) respectively, significantly higher than the value of 0.05°C of the control (BSA). Summarily, four homologous proteins showed good antifreeze property with the range from high to low as PicK>PicW2>PicW1>PicM. It suggested that they can be used as resources for genetic engineering of plant cold tolerance., (Copyright © 2017. Published by Elsevier B.V.)

Previously, the identification of fossil Pseudolarix at the species level has been based on the morphology of the bract-scale complex of the seed cone. The morphological consistence of fossils through most of the Cenozoic with extant P. amabilis has led them to be considered conspecific, suggesting that P. amabilis is an extraordinary example of morphological stasis. However, the lack of cuticular evidence, especially for the leaf-homologous bract, reduces the accuracy of fossil identification based on morphology, thus weakening the evidence for morphological stasis in P. amabilis. For the first time, we provide cuticular evidence of the bract-scale of fossil P. amabilis based on the bract-scale complex from the late Miocene Shengxian Formation, Zhejiang, East China, which improves the identification accuracy and reinforces the concept of morphological stasis in this species. Second, we preliminarily reveal the niche stability of P. amabilis, which corresponds to its morphological stasis. Finally, we infer that the late Miocene forest containing P. amabilis in Zhejiang was an evergreen sclerophyllous broad-leaved or mixed mesophytic forest, which combined with the evergreen broad-leaved forest suggested by previous megafossil studies, indicates the occurrence of vertical vegetation zonation.

In large parts of Europe, insecticide-free measures for protecting conifer plants are desired to suppress damage by the pine weevil Hylobius abietis (L.). Treatment with methyl jasmonate (MeJA), a chemical elicitor already used in crop production, may enhance expression of chemical defenses in seedlings in conifer regenerations. However, in a previous experiment, MeJA treatment resulted in substantially better field protection for Scots pine (Pinus sylvestris L.) than for Norway spruce (Picea abies (L.) Karst.). Hypothesizing that the variations may be at least due partly to volatiles released by MeJA-treated seedlings and their effects on pine weevil orientation, we examined tissue extracts of seedlings (from the same batches as previously used) by two-dimensional GC-MS. We found that the MeJA treatment increased contents of the monoterpene (-)-β-pinene in phloem (the weevil's main target tissue) of both tree species, however, the (-)-β-pinene/(-)-α-pinene ratio increased more in the phloem of P. sylvestris. We also tested the attractiveness of individual monoterpenes found in conifer tissues (needles and phloem) for pine weevils using an arena with traps baited with single-substance dispensers and pine twigs. Trap catches were reduced when the pine material was combined with a dispenser releasing (-)-β-pinene, (+)-3-carene, (-)-bornyl acetate or 1,8-cineole. However, (-)-α-pinene did not have this effect. Thus, the greater field protection of MeJA-treated P. sylvestris seedlings may be due to the selective induction of increases in contents of the deterrent (-)-β-pinene, in contrast to strong increases in both non-deterrent (-)-α-pinene and the deterrent (-)-β-pinene in P. abies seedlings.

Woodpeckers and other primary cavity excavators (PCEs) are important worldwide for excavating cavities in trees, and a large number of studies have examined their nesting preferences. However, quantitative measures of wood hardness have been omitted from most studies, and ecologists have focused on the effects of external tree- and habitat-level features on nesting. Moreover, information is lacking on the role of wood hardness in limiting nesting opportunities for this important guild. Here, we used an information theoretic approach to examine the role of wood hardness in multi-scale nest site selection and in limiting nesting opportunities for six species of North American PCEs. We found that interior wood hardness at nests (n = 259) differed from that at random sites, and all six species of PCE had nests with significantly softer interior wood than random trees (F1,517 = 106.15, P < 0.0001). Accordingly, interior wood hardness was the most influential factor in our models of nest site selection at both spatial scales that we examined: in the selection of trees within territories and in the selection of nest locations on trees. Moreover, regardless of hypothesized excavation abilities, all the species in our study appeared constrained by interior wood hardness, and only 4-14% of random sites were actually suitable for nesting. Our findings suggest that past studies that did not measure wood hardness counted many sites as available to PCEs when they were actually unsuitable, potentially biasing results. Moreover, by not accounting for nest site limitations in PCEs, managers may overestimate the amount of suitable habitat. We therefore urge ecologists to incorporate quantitative measures of wood hardness into PCE nest site selection studies, and to consider the limitations faced by avian cavity excavators in forest management decisions.

We developed models to describe the responses of four commonly examined leaf traits (mass per area, weight, area and nitrogen (N) concentration) to gradients of light, soil nutrients and tree height in three conifer species of contrasting shade tolerance. Our observational dataset from the sub-boreal spruce forests of British Columbia included subalpine fir (Abies lasioscarpa [Hook.] Nutt; high shade tolerance), interior spruce (Picea glauca × Picea engelmannii [Moench] Voss; intermediate shade tolerance) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia; low shade tolerance) saplings from 0.18 to 4.87 m tall, in 8-98% of total incident light, from field sites with <17.6 kg ha(-1) to >46.8 kg ha(-1) total dissolved N. Leaf weights and areas showed strong positive responses to light and height, but little or no response to soil nutrients. Parameter estimates indicated that the shape of leaf weight and area responses to light corresponded with shade tolerance ranking for the three species; pine had the most linear response whereas spruce and fir had asymptotic responses. Leaf N concentration responded positively to soil nutrients, negatively to light and idiosyncratically to height. The negative effect of light was only apparent on sites of high soil nutrient availability, and parameter estimates for the shape of the negative response also corresponded to shade tolerance ranking (apine = -0.79, aspruce = -0.15, afir = -0.07). Of the traits we measured, leaf mass per area showed the least response to light, soil nutrient and height gradients. Although it is a common practice in comparisons across many species, characterizing these conifers by mean values of their leaf traits would miss important intraspecific variation across environmental and size gradients. In these forests, parameter estimates representing the intraspecific variability of leaf trait responses can be used to understand relative shade tolerances., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Water stress is one of the primary selective forces in plant evolution. There are characters often cited as adaptations to water stress, but links between the function of these traits and adaptation to drying climates are tenuous. Here we combine distributional, climatic, and physiological evidence from 42 species of conifers to show that the evolution of drought resistance follows two distinct pathways, both involving the coordinated evolution of tissues regulating water supply (xylem) and water loss (stomatal pores) in leaves. Only species with very efficient stomatal closure, and hence low minimum rates of water loss, inhabit dry habitats, but species diverged in their apparent mechanism for maintaining closed stomata during drought. An ancestral mechanism found in Pinaceae and Araucariaceae species relies on high levels of the hormone abscisic acid (ABA) to close stomata during water stress. A second mechanism, found in the majority of Cupressaceae species, uses leaf desiccation rather than high ABA levels to close stomata during sustained water stress. Species in the latter group were characterized by xylem tissues with extreme resistance to embolism but low levels of foliar ABA after 30 d without water. The combination of low levels of ABA under stress with cavitation-resistant xylem enables these species to prolong stomatal opening during drought, potentially extending their photosynthetic activity between rainfall events. Our data demonstrate a surprising simplicity in the way conifers evolved to cope with water shortage, indicating a critical interaction between xylem and stomatal tissues during the process of evolution to dry climates.

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.

Climate change is anticipated to alter plant species distributions. Regional context, notably the spatial complexity of climatic gradients, may influence species migration potential. While high-elevation species may benefit from steep climate gradients in mountain regions, their persistence may be threatened by limited suitable habitat as land area decreases with elevation. To untangle these apparently contradictory predictions for mountainous regions, we evaluated the climatic suitability of four coniferous forest tree species of the western United States based on species distribution modeling (SDM) and examined changes in climatically suitable areas under predicted climate change. We used forest structural information relating to tree species dominance, productivity, and demography from an extensive forest inventory system to assess the strength of inferences made with a SDM approach. We found that tree species dominance, productivity, and recruitment were highest where climatic suitability (i.e., probability of species occurrence under certain climate conditions) was high, supporting the use of predicted climatic suitability in examining species risk to climate change. By predicting changes in climatic suitability over the next century, we found that climatic suitability will likely decline, both in areas currently occupied by each tree species and in nearby unoccupied areas to which species might migrate in the future. These trends were most dramatic for high elevation species. Climatic changes predicted over the next century will dramatically reduce climatically suitable areas for high-elevation tree species while a lower elevation species, Pinus ponderosa, will be well positioned to shift upslope across the region. Reductions in suitable area for high-elevation species imply that even unlimited migration would be insufficient to offset predicted habitat loss, underscoring the vulnerability of these high-elevation species to climatic changes., (© 2013 John Wiley & Sons Ltd.)

Background: There is a rapidly growing awareness that plant peptide signalling molecules are numerous and varied and they are known to play fundamental roles in angiosperm plant growth and development. Two closely related peptide signalling molecule families are the CLAVATA3-EMBRYO-SURROUNDING REGION (CLE) and CLE-LIKE (CLEL) genes, which encode precursors of secreted peptide ligands that have roles in meristem maintenance and root gravitropism. Progress in peptide signalling molecule research in gymnosperms has lagged behind that of angiosperms. We therefore sought to identify CLE and CLEL genes in gymnosperms and conduct a comparative analysis of these gene families with angiosperms., Results: We undertook a meta-analysis of the GenBank/EMBL/DDBJ gymnosperm EST database and the Picea abies and P. glauca genomes and identified 93 putative CLE genes and 11 CLEL genes among eight Pinophyta species, in the genera Cryptomeria, Pinus and Picea. The predicted conifer CLE and CLEL protein sequences had close phylogenetic relationships with their homologues in Arabidopsis. Notably, perfect conservation of the active CLE dodecapeptide in presumed orthologues of the Arabidopsis CLE41/44-TRACHEARY ELEMENT DIFFERENTIATION (TDIF) protein, an inhibitor of tracheary element (xylem) differentiation, was seen in all eight conifer species. We cloned the Pinus radiata CLE41/44-TDIF orthologues. These genes were preferentially expressed in phloem in planta as expected, but unexpectedly, also in differentiating tracheary element (TE) cultures. Surprisingly, transcript abundances of these TE differentiation-inhibitors sharply increased during early TE differentiation, suggesting that some cells differentiate into phloem cells in addition to TEs in these cultures. Applied CLE13 and CLE41/44 peptides inhibited root elongation in Pinus radiata seedlings. We show evidence that two CLEL genes are alternatively spliced via 3'-terminal acceptor exons encoding separate CLEL peptides., Conclusions: The CLE and CLEL genes are found in conifers and they exhibit at least as much sequence diversity in these species as they do in other plant species. Only one CLE peptide sequence has been 100% conserved between gymnosperms and angiosperms over 300 million years of evolutionary history, the CLE41/44-TDIF peptide and its likely conifer orthologues. The preferential expression of these vascular development-regulating genes in phloem in conifers, as they are in dicot species, suggests close parallels in the regulation of secondary growth and wood formation in gymnosperm and dicot plants. Based on our bioinformatic analysis, we predict a novel mechanism of regulation of the expression of several conifer CLEL peptides, via alternative splicing resulting in the selection of alternative C-terminal exons encoding separate CLEL peptides.

Premise of the Study: Seed cone scales assigned to the genus Schizolepidopsis are widespread in Late Triassic to Cretaceous Eurasian deposits. They have been linked to the conifer family Pinaceae based on associated vegetative remains, but their exact affinities are uncertain. Recently discovered material from the Early Cretaceous of Mongolia reveals important new information concerning Schizolepidopsis cone scales and seeds, and provides support for a relationship between the genus and extant Pinaceae., Methods: Specimens were collected from Early Cretaceous (probable Aptian-Albian) lignite deposits in central Mongolia. Lignite samples were disaggregated, cleaned in hydrofluoric acid, and washed in water. Specimens were selected for further study using light and electron microscopy., Key Results: Schizolepidopsis canicularis seed cones consist of loosely arranged, bilobed ovulate scales subtended by a small bract. A single inverted seed with an elongate micropyle is borne on each lobe of the ovulate scale. Each seed has a wing formed by the separation of the adaxial surface of the ovulate scale., Conclusions: Schizolepidopsis canicularis produced winged seeds that formed in a manner that is unique to Pinaceae among extant conifers. We do not definitively place this species in Pinaceae pending more complete information concerning its pollen cones and vegetative remains. Nevertheless, this material suggests that Schizolepidopsis may be important for understanding the early evolution of Pinaceae, and may potentially help reconcile the appearance of the family in the fossil record with results based on phylogenetic analyses of molecular data.

Forests around the world are subject to risk of high rates of tree growth decline and increased tree mortality from combinations of climate warming and drought, notably in semi-arid settings. Here, we assess how climate warming has affected tree growth in one of the world's most extensive zones of semi-arid forests, in Inner Asia, a region where lack of data limits our understanding of how climate change may impact forests. We show that pervasive tree growth declines since 1994 in Inner Asia have been confined to semi-arid forests, where growing season water stress has been rising due to warming-induced increases in atmospheric moisture demand. A causal link between increasing drought and declining growth at semi-arid sites is corroborated by correlation analyses comparing annual climate data to records of tree-ring widths. These ring-width records tend to be substantially more sensitive to drought variability at semi-arid sites than at semi-humid sites. Fire occurrence and insect/pathogen attacks have increased in tandem with the most recent (2007-2009) documented episode of tree mortality. If warming in Inner Asia continues, further increases in forest stress and tree mortality could be expected, potentially driving the eventual regional loss of current semi-arid forests., (© 2013 John Wiley & Sons Ltd.)

Predicting climate-driven changes in plant distribution is crucial for biodiversity conservation and management under recent climate change. Climate warming is expected to induce movement of species upslope and towards higher latitudes. However, the mechanisms and physiological processes behind the altitudinal and latitudinal distribution range of a tree species are complex and depend on each tree species features and vary over ontogenetic stages. We investigated the altitudinal distribution differences between juvenile and adult individuals of seven major European tree species along elevational transects covering a wide latitudinal range from southern Spain (37°N) to northern Sweden (67°N). By comparing juvenile and adult distributions (shifts on the optimum position and the range limits) we assessed the response of species to present climate conditions in relation to previous conditions that prevailed when adults were established. Mean temperature increased by 0.86 °C on average at our sites during the last decade compared with previous 30-year period. Only one of the species studied, Abies alba, matched the expected predictions under the observed warming, with a maximum abundance of juveniles at higher altitudes than adults. Three species, Fagus sylvatica, Picea abies and Pinus sylvestris, showed an opposite pattern while for other three species, such as Quercus ilex, Acer pseudoplatanus and Q. petraea, we were no able to detect changes in distribution. These findings are in contrast with theoretical predictions and show that tree responses to climate change are complex and are obscured not only by other environmental factors but also by internal processes related to ontogeny and demography., (© 2013 John Wiley & Sons Ltd.)

The present study investigated the genetic diversity, population structure, F ST outliers, and extent and pattern of linkage disequilibrium in five populations of Keteleeria davidiana var. formosana, which is listed as a critically endangered species by the Council of Agriculture, Taiwan. Twelve amplified fragment length polymorphism primer pairs generated a total of 465 markers, of which 83.74% on average were polymorphic across populations, with a mean Nei's genetic diversity of 0.233 and a low level of genetic differentiation (approximately 6%) based on the total dataset. Linkage disequilibrium and HICKORY analyses suggested recent population bottlenecks and inbreeding in K. davidiana var. formosana. Both STRUCTURE and BAPS observed extensive admixture of individual genotypes among populations based on the total dataset in various clustering scenarios, which probably resulted from incomplete lineage sorting of ancestral variation rather than a high rate of recent gene flow. Our results based on outlier analysis revealed generally high levels of genetic differentiation and suggest that divergent selection arising from environmental variation has been driven by differences in temperature, precipitation, and humidity. Identification of ecologically associated outliers among environmentally disparate populations further support divergent selection and potential local adaptation.

Stomata control tree transpiration by sensing and integrating environmental signals originating in the atmosphere and soil, and co-occurring species may differ in inherent stomatal sensitivity to these above- and belowground signals and in the types of signals to which they respond. Stomatal responsiveness to environmental signals is likely to differ across species having different types of wood (e.g., ring-porous, diffuse-porous and coniferous) because each wood type differs in the structure, size and spatial distribution of its xylem conduits as well as in the scaling of hydraulic properties with stem diameter. The objective of this study was to evaluate the impact of variation in soil water availability and atmospheric evaporative demand on stomatal regulation of transpiration in seven co-occurring temperate deciduous forest species representing three wood types. We measured whole-tree sap flux and soil and atmospheric variables in a mixed deciduous forest in central Pennsylvania over the course of a growing season characterized by severe drought and large fluctuations in atmospheric vapor pressure deficit (D). The relative sensitivity of sap flux to soil drying was ∼2.2-2.3 times greater in the diffuse-porous and coniferous species than in the ring-porous species. Stomata of the ring-porous oaks were only about half as responsive to increased D as those of trees of the other two wood types. These differences in responsiveness to changes in the below- and aboveground environment implied that regulation of leaf water potential in the ring-porous oaks was less stringent than that in the diffuse-porous angiosperms or the conifers. The results suggest that increases in the frequency or intensity of summer droughts in the study region could have multiple consequences for forest function, including altered successional time courses or climax species composition and cumulative effects on whole-tree architecture, resulting in a structural and physiological legacy that would restrict the ability of trees to respond rapidly to more favorable growth conditions.

Insect herbivores are integral to terrestrial ecosystems. They provide essential food for higher trophic levels and aid in nutrient cycling. In general, research tends to relate individual insect herbivore species to host plant identity, where a species will show preference for one host over another. In contrast, insect herbivore assemblages are often related to host plant richness where an area with a higher richness of hosts will also have a higher richness of herbivores. In this study, the ability of these two approaches (host plant identity/abundance vs. host plant richness) to describe the diversity, richness, and abundance of an herbivorous Lepidoptera assemblage in temperate forest fragments in southern Canada is tested. Analyses indicated that caterpillar diversity, richness, and abundance were better described by quadrat-scale host plant identity and abundance than by host plant richness. Most host plant-herbivore studies to date have only considered investigating host plant preferences at a species level; the type of assemblage level preference shown in this study has been rarely considered. In addition, host plant replacement simulations indicate that increasing the abundance of preferred host plants could increase Lepidoptera richness and abundance by as much as 30% and 40% respectively in disturbed remnant forest fragments. This differs from traditional thinking that suggests higher levels of insect richness can be best obtained by maximizing plant richness. Host plant species that are highly preferred by the forest-dwelling caterpillar assemblage should be given special management and conservation considerations to maximize biodiversity in forest communities.

Dehydrins are thought to play an essential role in the response, acclimation and tolerance to different abiotic stresses, such as cold and drought. These proteins have been classified into five groups according to the presence of conserved and repeated motifs in their amino acid sequence. Due to their putative functions in the response to stress, dehydrins have been often used as candidate genes in studies on population variability and local adaptation to environmental conditions. However, little is still known regarding the differential role played by such groups or the mechanism underlying their function. Based on the sequences corresponding to dehydrins available in public databases we have isolated eight different dehydrins from cDNA of Pinus pinaster. We have obtained also their genomic sequences and identified their intron/exon structure. Quantitative RT-PCR analysis of their expression pattern in needles, stems and roots during a severe and prolonged drought stress, similar to the ones trees must face in nature, is also reported. Additionally, we have identified two amino acid motifs highly conserved and repeated in Pinaceae dehydrins and absent in angiosperms, presumably related to the divergent expression profiles observed.

Tricholoma matsutake produces commercially valuable, yet uncultivable, mushrooms (matsutake) in association with pines in the Far East and Scandinavia and with both pines and oaks in the foothills of Tibet. Other matsutake mushrooms, such as Tricholoma anatolicum from the Mediterranean regions and Tricholoma magnivelare and Tricholoma sp. from the North Pacific Coast area of Canada and North America as well as Mexico, respectively, are associated with pines or oaks in their natural habitats. Tricholoma bakamatsutake and Tricholoma fulvocastaneum from Asia produce moderately valuable matsutake mushrooms and are solely associated with Fagaceae in nature. In this study, we demonstrate for the first time that matsutake mushrooms from Scandinavia, Mediterranean regions, North America, and Tibet form ectomycorrhizae with Pinus densiflora similar to the Far East T. matsutake. In general, worldwide T. matsutake and the symbionts of Pinaceae colonize the rhizospheres of P. densiflora as well as T. matsutake isolated from the host plant. However, T. fulvocastaneum and T. bakamatsutake formed a discontinuous Hartig net and no Hartig net, respectively, and colonized to a lesser extent as compared to T. matsutake. The data suggest that conifer-associated matsutake mushrooms in their native habitat will associate symbiotically with the Asian red pine.

Why particular invasions succeed and others fail is not well understood. The role of soil biota has been proposed as important. However, the role of mutualists has received much less attention than that of pathogens. Here we report that lack of adequate ectomycorrhizal fungi hinders invasion by exotic Pinaceae on Isla Victoria, Argentina, by reducing both the probability of establishment and growth of invading individuals. More than one hundred exotic tree species were introduced to this island ca. 80 years ago, but invasive trees are found in high densities only in areas adjacent to plantations. With a series of greenhouse and field experiments we found lower mycorrhizal colonization levels and few fungal species far from original plantings, and key fungal mutualists are confined to areas near plantations, probably owing to dispersal limitations. Low inoculum levels far from the plantations are retarding the invasion. Our experiments indicate that positive interactions belowground can play a key but underappreciated role in invasion dynamics.

Adequate leaf hydraulic conductance (Kleaf) is critical for preventing transpiration-induced desiccation and subsequent stomatal closure that would restrict carbon gain. A few studies have reported midday depression of Kleaf (or petiole conductivity) and its subsequent recovery in situ, but the extent to which this phenomenon is universal is not known. The objectives of this study were to measure Kleaf, using a rehydration kinetics method, (1) in the laboratory (under controlled conditions) across a range of water potentials to construct vulnerability curves (VC) and (2) over the course of the day in the field along with leaf water potential and stomatal conductance (gs). Two broadleaf (one evergreen, Arbutus menziesii Pursh., and one deciduous, Quercus garryana Dougl.) and two coniferous species (Pinus ponderosa Dougl. and Pseudotsuga menziesii [Mirbel]) were chosen as representative of different plant types. In addition, Kleaf in the laboratory and leaf water potential in the field were measured for three tropical evergreen species (Protium panamense (Rose), Tachigalia versicolor Standley and L.O. Williams and Vochysia ferruginea Mart) to predict their daily changes in field Kleaf in situ. It was hypothesized that in the field, leaves would close their stomata at water potential thresholds at which Kleaf begins to decline sharply in laboratory-generated VC, thus preventing substantial losses of Kleaf. The temperate species showed a 15-66% decline in Kleaf by midday, before stomatal closure. Although there were substantial midday declines in Kleaf, recovery was nearly complete by late afternoon. Stomatal conductance began to decrease in Pseudotsuga, Pinus and Quercus once Kleaf began to decline; however, there was no detectable reduction in gs in Arbutus. Predicted Kleaf in the tropical species, based on laboratory-generated VC, decreased by 74% of maximum Kleaf in Tachigalia, but only 22-32% in Vochysia and Protium. The results presented here, from the previous work of the authors and from other published studies, were consistent with two different strategies regarding daily maintenance of Kleaf: (1) substantial loss and subsequent recovery or (2) a more conservative strategy of loss avoidance.

To provide baseline data for physiological studies of extreme low-temperature (LT) tolerance in boreal conifers, we profiled LT stress responses, liquid nitrogen (LN(2))-quench tolerance, and sugar concentrations in foliage of boreal-temperate species pairs in the genera Abies, Picea and Pinus, growing in an arboretum in a temperate oceanic climate from August 2006 through April 2007. The boreal species acclimated more rapidly and deeply than the temperate species, acquiring LN(2)-quench tolerance by late November, despite unusually warm conditions throughout the autumn and early winter. Maximum LT tolerance in the temperate species was in the -25 to -35 degrees C range, and was reached only after a period of freezing temperatures in late January and February. During LT acclimation in the temperate species, sigmoid temperature-relative electrolyte leakage (REL) curves shifted toward lower temperatures, whereas in boreal species there was both a temperature shift and a lowering of the maximum REL until it fell below a threshold associated with irreversible injury. These differences may reflect differences in mechanisms of LT acclimation and LT tolerance. The concentrations of total and individual sugars did not show a clear pattern that could differentiate the boreal and temperate groups. Raffinose and, in three of the six species, stachyose showed the closest association with LT tolerance. Sugar concentrations, principally sucrose, decreased during mild weather, perhaps because of respiratory losses or phloem export, and increased after periods of freezing temperatures. Low-temperature acclimation in boreal species appears to follow a rigid program that may affect their ability to avoid excessive respiratory losses in the event of continued climate warming in boreal regions.

Eighty-four mature Norway spruce (Picea abies L. Karst), silver fir (Abies alba Mill) and Scots pine (Pinus sylvestris L.) trees were winched over to determine the maximum resistive turning moment (M(a)) of the root-soil system, the root-soil plate geometry, the azimuthal orientation of root growth, and the occurrence of root rot. The calculation of M(a), based on digital image tracking of stem deflection, accounted not only for the force application and its changing geometry, but also for the weight of the overhanging tree, representing up to 42% of M(a). Root rot reduced M(a) significantly and was detected in 25% of the Norway spruce and 5% of the silver fir trees. Excluding trees with root rot, differences in M(a) between species were small and insignificant. About 75% of the variance in M(a) could be explained by one of the four variables--tree mass, stem mass, stem diameter at breast height squared times tree height, and stem diameter at breast height squared. Among the seven allometric variables assessed above ground, stem diameter at breast height best described the root-soil plate dimensions, but the correlations were weak and the differences between species were insignificant. The shape of the root-soil plate was well described by a depth-dependent taper model with an elliptical cross section. Roots displayed a preferred azimuthal orientation of growth in the axis of prevailing winds, and the direction of frequent weak winds matched the orientation of growth better than that of rare strong winds. The lack of difference in anchorage parameters among species probably reflects the similar belowground growth conditions of the mature trees.

Variation in tree stem diameter results from reversible shrinking and swelling and irreversible radial growth, all processes that are influenced by tree water status. To assess the causal effects of water and temperature on stem radial variation (DeltaR) and maximum daily shrinkage (MDS), the diurnal cycle was divided into three phases: contraction, expansion and stem radius increment. Diurnal cycles were measured during 1996-2004 in Picea abies (L.) Karst., Pinus cembra L. and Larix decidua Mill. in a timberline ecotone to understand the links between stem diameter variation (v; defined as MDS or DR), phase duration (h), and weather or sap flow descriptors (d). We demonstrated that a high proportion of MDS and DeltaR was explained by h because of the nonlinearity of the physiological responses to weather d. By causal modeling, we tested whether the relationship between d and v was due to h (lack of causal relationship between d and v) or to both d and h (double cause). The results of this modeling added new physiological insight into daily growth-climate relationships. Negative correlations were found between DeltaR and air temperature owing to the negative effect of temperature on h only, and did not correspond to a direct effect on tree growth mediated by an alteration in metabolic activities. Precipitation had two main effects: a direct effect on DeltaR and an indirect effect mediated through an effect on h. A reduction in sap flow at night led to an increase in DeltaR for P. abies and L. decidua, but not for P. cembra.

To advance our understanding of host effects on the community structure of ectomycorrhizal fungi (EMF), EMF communities were compared among different host species, genera and families in two mixed conifer-broadleaf forests in Japan. Using molecular identification methods we examined EMF root tips of eight coexisting species belonging to six genera (three families): Abies and Tsuga (Pinaceae), Betula and Carpinus (Betulaceae) and Fagus and Quercus (Fagaceae). In total, 205 EMF species were detected, and the total richness was estimated to exceed 300 species using major estimators. Of the 55 EMF species occurring three or more times, eight showed significantly biased host preference. A Mantel test showed a significantly negative correlation between EMF community similarity and host taxonomic distance. Detrended correspondence analysis separated EMF communities mainly by host taxonomic and successional status. Thus, EMF communities are similar on hosts with similar taxonomic and successional status. A significant proportion of EMF exhibited host specificity, which may contribute to the extremely diverse EMF community in conifer-broadleaf forests.

Drought stress can cause xylem embolism in trees when the water potential (psi) in the xylem falls below specific vulnerability thresholds. At the alpine timberline, frost drought is known to cause excessive winter embolism unless xylem vulnerability or transpiration is sufficiently reduced to avoid critical psi. We compared annual courses of psi and embolism in Picea abies, Pinus cembra, Pinus mugo, Larix decidua, and Juniperus communis growing at the timberline vs. low altitude. In addition, vulnerability properties and related anatomical parameters as well as wood density (D(t)) and wall reinforcement (wall thickness related to conduit diameter) were studied. This allowed an estimate of stress intensities as well as a detection of adaptations that reduce embolism formation. At the alpine timberline, psi was lowest during winter with corresponding embolism rates of up to 100% in three of the conifers studied. Only Pinus cembra and Larix decidua avoided winter embolism due to moderate psi. Minor embolism was observed at low altitude where the water potentials of all species remained within a narrow range throughout the year. Within species, differences in psi50 (psi at 50% loss of conductivity) at high vs. low altitude were less than 1 MPa. In Picea abies and Pinus cembra, psi50 was more negative at the timberline while, in the other conifer species, psi50 was more negative at low altitude. Juniperus communis exhibited the lowest (-6.4 +/- 0.04 MPa; mean +/- SE) and Pinus mugo the highest psi50 (-3.34 +/- 0.03 MPa). In some cases, D(t) and tracheid wall reinforcement were higher than in previously established relationships of these parameters with psi50, possibly because of mechanical demands associated with the specific growing conditions. Conifers growing at the alpine timberline were exposed to higher drought stress intensities than individuals at low altitude. Frost drought during winter caused high embolism rates which were probably amplified by freeze-thaw stress. Although frost drought had a large effect on plant water transport, adaptations in hydraulic safety and related anatomical parameters were observed in only a few of the conifer species studied.

As trees grow tall and the resistance of the hydraulic pathway increases, water supply to foliage may decrease forcing stomata to close and CO2 uptake to decline. Several structural (e.g. biomass allocation) and physiological adjustments, however, may partially or fully compensate for such hydraulic constraints and prevent limitations on CO2 uptake and growth. The degree to which trees compensate for hydraulic constraints as they grow tall may depend on the costs and benefits associated with hydraulic compensation according to their ecology and life history. Because later successional Rocky Mountain conifers are more shade tolerant, optimization of CO2 uptake as trees grow tall and shade increases may confer greater benefits than in earlier successional species. If so, higher compensation for hydraulic constraints is expected in later successional species relative to co-occurring earlier successional species. I have examined height-related changes of crown stomatal conductance on a leaf area basis (G(LA)) and leaf to sapwood ratios (A(L):A(S)) for five conifer species in the northern Rocky Mountains. Species were arranged in pairs, each pair consisting of an early and late successional species. For high elevations I used, respectively, whitebark pine (Pinus albicaulis) and subalpine fir (Abies lasiocarpa); for mid-elevations, western larch (Larix occidentalis) and Douglas-fir (Pseudotsuga menziesii); for lower elevations, ponderosa pine (Pinus ponderosa) and Douglas-fir. A(L):A(S) either decreased (subalpine fir, ponderosa pine), remained constant (Douglas-fir, western larch) or increased (whitebark pine) with tree height. As hypothesized, earlier successional species (ponderosa pine, whitebark pine and western larch) exhibited significantly stronger decreases of G(LA) with tree height relative to their later successional pairs (Douglas-fir and subalpine fir), which fully compensated for height-related hydraulic constraints on G(LA). A life history approach that takes into account the optimization of size- and species-specific ecological functions may also help researchers better understand biomass allocation and hydraulic function in trees.

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.

Simultaneous saccharification and fermentation (SSF) of wood has been modeled for the past 15-20 years, but the substrates used for model evaluation have so far not included pretreated softwood. In the present study, data from lab-scale batch SSF of SO2-impregnated, steam-pretreated spruce chips were used to evaluate a model found in the literature. The model, which was somewhat modified, consists of a number of nonlinear, coupled ordinary differential equations, which were solved numerically. Some parameter values were fitted to data by use of least-squares minimization. A difficulty in parameter estimation was the lack of cellobiose measurements, something that was relieved by adding assumptions about parameter relations. The simulated concentration profiles agreed well with the measured concentrations of glucose and ethanol. It is therefore concluded that the basic model features apply to softwood SSF. The model predicts rate saturation with respect to enzyme concentration at concentrations above 60 FPU/g cellulose, although this was not observed in the experimental data, which only comprised enzyme concentrations up to 32 FPU/g cellulose.

Type and dynamics of age structure of Keteleeria davidiana var. chien-peii population and its relationship to community succession and environment were analyzed. The results showed that the age structures were classified into four types:growing, stable, initial and middle sensecent types. The survival curves had concave, convex, disconnected and dotted shapes. Along with development and succession of community, the age structure changes from growing, stabilized, senescent to residual type. The important factors causing change to the age structure were biological features of Keteleeria davidiana var. chien-peii, development of broad-leaved tree species, geographic isolation, and human disturbance.

Twenty-four temperate tree species were classified into three groups based on cluster analysis of relative growth rate, nitrogen concentration, nitrogen-production efficiency, nitrogen-distribution ratio and nitrogen-use efficiency as follows: Group I (Asteridae and Rosidae), Group II (Dilleniidae and Hamamelidae) and Group III (Coniferopsidae). Relative growth rate (RGR) was high in Group II, moderate in Group I and low in Group III. The regression coefficient for the relationship between RGR and leaf nitrogen concentration was higher in Group II than in Group I, and no relationship was observed in Group III. Parameter analysis of RGR indicated that RGR per unit leaf nitrogen was important for all three groups, but that the allocation of nitrogen to leaves was particularly important in Groups I and II. The ratio of dark respiratory rate (R) to net photosynthetic rate (A) was higher in Group I than in Group II. Neither A nor R was measured in the Group III species. A linear relationship was observed between leaf nitrogen concentration and A in Group II, but this relationship was not evident in Group I.

The competition among trees within a stand includes aboveground and underground parts. How to determine the zone of competition influence is important for predicting tree growth. We divide the zone into two parts: one part is defined as inside radius, the distance from nearest competitors to subject tree, in which, the competition influence is the sum of both aboveground and underground; the another part is defined as outside radius, the distance of competitors to subject tree, where the competition influence comes mostly from underground. Thus, the competition index, based upon the divided two radii, could be decomposed into different components. We used the data of manchurian ash(MA) (Fraxinus mandshurica) and dahurian larch(DL) (Larix gmelini) from pure and mixed stands to analyze the competition influence of aboveground and underground in inside and of underground in outside radius. The results show that for MA in pure stand, the competition effect was 59.06% from inside radius, and 40.94% from outside radius. For DL in pure stand, the competition effects in both radii were 71.31% and 28.69%, respectively. There was a stronger competition influence in inside than in outside radius. Underground competitions were the important issues for both species in pure stands. In a mixed stand of MA and DL, there were two species in both radii, and the effects of competition came from intraspecific and interspecific competitors. For subject tree of MA, the effect of interspecific competition was 49.55%, and that of intraspecific was 50.45%. For subject tree of DL, the effects were 81.11% and 18.89%, respectively. DL displayed a stronger interspecific competition than MA in the zone of competition influence, and was a main effect of competition on MA. For subject tree of MA, the effect of competition from inside radius was 25.33%, which was the sum of competition influence from aboveground and underground, and about 74.67% was from outside radius. For subject tree of DL, both in inside and outside radii were 55.37% and 44.64%, respectively. Comparing with the information from both species and two radii, the root-root competition from underground was an important component. DL grew well, and could reduce the impact of intraspecific competition for MA when both species mixed in a stand.

Based on distance-dependent competition models and neighborhood interference models, the competition index components and competition effect coefficients were used to quantify the spatial competition among trees in the mixed plantation of ash(Fraxinus mandshurica) and larch(Larix gmelini). Taking each component of competition index as independent variables and the yearly volume growth as dependent variable, the competition-growth model was established with good regression result. The study provided a new approach to deal with the further improvement of spatial competition models and the plant competition-growth relationship ecological study.