Extreme low temperature tolerance in woody plants

Woody plants in boreal to arctic environments and high mountains survive prolonged exposure to temperatures below -40°C and minimum temperatures below -60°C, and laboratory tests show that many of these species can also survive immersion in liquid nitrogen at -196°C. Studies of biochemical changes that occur during acclimation, including recent proteomic and metabolomic studies, have identified changes in carbohydrate and compatible solute concentrations, membrane lipid composition, and proteins, notably dehydrins, that may have important roles in survival at extreme low temperature (ELT). Consideration of the biophysical mechanisms of membrane stress and strain lead to the following hypotheses for cellular and molecular mechanisms of survival at ELT: (1) Changes in lipid composition stabilize membranes at temperatures above the lipid phase transition temperature (-20 to -30°C), preventing phase changes that result in irreversible injury. (2) High concentrations of oligosaccharides promote vitrification or high viscosity in the cytoplasm in freeze-dehydrated cells, which would prevent deleterious interactions between membranes. (3) Dehydrins bind membranes and further promote vitrification or act stearically to prevent membrane–membrane interactions.© 2015 Strimbeck, Schaberg, Fossdal, Schröder and Kjellsen. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms

The promise and peril of intensive-site-based ecological research: insights from the Hubbard Brook ecosystem study

Abstract. Ecological research is increasingly concentrated at particular locations or sites. This trend reflects a variety of advantages of intensive, site-based research, but also raises important questions about the nature of such spatially delimited research: how well does site based research represent broader areas, and does it constrain scientific discovery?We provide an overview of these issues with a particular focus on one prominent intensive research site: the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA. Among the key features of intensive sites are: long-term, archived data sets that provide a context for new discoveries and the elucidation of ecological mechanisms; the capacity to constrain inputs and parameters, and to validate models of complex ecological processes; and the intellectual cross-fertilization among disciplines in ecological and environmental sciences. The feasibility of scaling up ecological observations from intensive sites depends upon both the phenomenon of interest and the characteristics of the site. An evaluation of deviation metrics for the HBEF illustrates that, in some respects, including sensitivity and recovery of streams and trees from acid deposition, this site is representative of the Northern Forest region, of which HBEF is a part. However, the mountainous terrain and lack of significant agricultural legacy make the HBEF among the least disturbed sites in the Northern Forest region. Its relatively cool, wet climate contributes to high stream flow compared to other sites. These similarities and differences between the HBEF and the region can profoundly influence ecological patterns and processes and potentially limit the generality of observations at this and other intensive sites. Indeed, the difficulty of scaling up may be greatest for ecological phenomena that are sensitive to historical disturbance and that exhibit the greatest spatiotemporal variation, such as denitrification in soils and the dynamics of bird communities. Our research shows that end member sites for some processes often provide important insights into the behavior of inherently heterogeneous ecological processes. In the current era of rapid environmental and biological change, key ecological responses at intensive sites will reflect both specific local drivers and regional trends

Age, allocation and availability of nonstructural carbon in mature red maple trees

The allocation of nonstructural carbon (NSC) to growth, metabolism and storage remains poorly understood, but is critical for the prediction of stress tolerance and mortality. We used the radiocarbon (14C) ‘bomb spike’ as a tracer of substrate and age of carbon in stemwood NSC, CO2 emitted by stems, tree ring cellulose and stump sprouts regenerated following harvesting in mature red maple trees. We addressed the following questions: which factors influence the age of stemwood NSC?; to what extent is stored vs new NSC used for metabolism and growth?; and, is older, stored NSC available for use? The mean age of extracted stemwood NSC was 10 yr. More vigorous trees had both larger and younger stemwood NSC pools. NSC used to support metabolism (stem CO2) was 1–2 yr old in spring before leaves emerged, but reflected current-year photosynthetic products in late summer. The tree ring cellulose 14C age was 0.9 yr older than direct ring counts. Stump sprouts were formed from NSC up to 17 yr old. Thus, younger NSC is preferentially used for growth and day-to-day metabolic demands. More recently stored NSC contributes to annual ring growth and metabolism in the dormant season, yet decade-old and older NSC is accessible for regrowth

Modeling the impacts of hemlock woolly adelgid infestation and presalvage harvesting on carbon stocks in northern hemlock forests

To better understand the potential impact of HWA and presalvage activities on carbon (C) dynamics in northern hemlock stands, we used the Forest Vegetation Simulator and Forest Inventory Analysis data to model C storage and successional pathways under four scenarios: presalvage harvesting; HWA-induced mortality; presalvage harvesting plus HWA-induced mortality, and a no disturbance control. Our simulation showed that all treatments differed in total C storage in the short term, with HWA-induced mortality providing the highest total C storage due to regeneration and ingrowth of replacement species combined with retention of standing and downed dead wood. At the end of the 150-year simulation, all disturbance scenarios had significantly lower total C than the control. The cumulative net C gain, was lower for the two presalvage than HWA scenario, indicating that allowing HWA to progress naturally through a stand may result in the least impact to long-term C sequestration and net C storage. While differences were not significant on low hemlock density stands, impacts to the estimated 267,000 hectares of northeastern forests where hemlock is dominant could result in conversion to red maple and a net loss of over 4 million metric tons of potentially sequestered C over the next 150 years.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

70 years of forest growth and community dynamics in an undisturbed northern hardwood forest

Long-term forest inventories provide a unique opportunity to quantify changes in forest structure and evaluate how changes compare to current stand development models. An examination of a 70-year record at the Bartlett Experimental Forest, NH indicated that while species abundances have primarily changed as expected under natural succession, some unexpected results were also detected. This included a significant decline in sugar maple (Acer saccharum) abundance driven by reduced regeneration, and increases in red spruce (Picea rubens) at the expense of sympatric balsam fir (Abies balsamea) and hardwoods at upper elevations. In contrast with accepted stand development models, biomass continues to accrue on these mid- to late-successional forests. Importantly, biomass accumulated at even greater rates in recent decades compared to historical norms. These results support evidence that the anthropogenic influences of a changing climate and the legacy of acid deposition may be altering stand dynamics in northeastern forests.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

Comparative growth trends of five northern hardwood and montane tree species reveal divergent trajectories and response to climate

In the northeastern US, tree declines associated with acid deposition-induced calcium depletion have been documented, notably for red spruce (Picea rubens Sarg.) and sugar maple (Acer saccharum Marsh.). There is conflicting evidence if co-occurring tree species have capitalized on these declines or suffered similar growth reductions; and, how growth has fluctuated relative to environmental variables. We examined five species along three elevational transects on Mt. Mansfield, Vermont: sugar maple, red spruce, red maple (Acer rubrum L.), yellow birch (Betula alleghaniensis, Britton), and balsam fir (Abies balsamea, [L.] Mill.). We found baseline differences in growth. Red maple and yellow birch had the highest growth, sugar maple and red spruce intermediate, and balsam fir the lowest. While some of year-to-year declines were associated with specific stress events, protracted patterns, such as recent increases in red spruce and red maple growth, were correlated with increased temperature and cooling degree days (heat index). For most species and elevations, there was a positive association between temperature and growth, but a negative association with growth the following year. Based on our comparisons, for some species growth at Mt. Mansfield aligns with regional trends and suggests that patterns assessed here may be indicative of the broader region.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

Experimental branch cooling increases foliar sugar and anthocyanin concentrations in sugar maple at the end of the growing season

Autumnal leaf anthocyanin expression is enhanced following exposure to a variety of environmental stresses and may be of adaptive benefit protecting leaves from those stresses, thereby allowing for prolonged sugar and nutrient resorption. Past work has shown that experimentally-induced sugar accumulations following branch girdling triggers anthocyanin biosynthesis. We hypothesized that reduced phloem transport at low autumnal temperatures may increase leaf sugar concentrations that stimulate anthocyanin production, resulting in enhanced tree- and landscape-scale color change. We used refrigerant-filled tubing to cool individual branches in a mature sugar maple (Acer saccharum Marsh) tree to test whether phloem cooling would trigger foliar sugar accumulations and enhance anthocyanin biosynthesis. Cooling increased foliar sucrose, glucose, and fructose concentrations 2- to nearly 10-fold (depending on the specific sugar and sampling date) relative to controls, and increased anthocyanin concentrations by approximately the same amount. Correlation analyses indicated a strong and steady positive relationship between anthocyanin and sugar concentrations, which was consistent with a mechanistic link between cooling-induced changes in these constituents. Tested here at the branch-level, we propose that low temperature-induced reductions in phloem transport may be responsible for increases in foliar sugars that trigger anthocyanin displays at grander scales.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

Anthropogenic alterations of genetic diversity within tree populations: Implications for forest ecosystem resilience

Healthy forests provide many of the essential ecosystem services upon which all life depends. Genetic diversity is an essential component of long-term forest health because it provides a basis for adaptation and resilience to environmental stress and change. In addition to natural processes, numerous anthropogenic factors deplete forest genetic resources. Genetic losses could be particularly consequential now because robust resilience is needed to respond to a growing number, variety, and frequency of stress exposures. Silvicultural management that selectively removes trees (and their genes) from forests may be another force reshaping forest gene pools. Although data concerning the influence of silvicultural management on genetic resources in temperate forests is somewhat mixed, through the genetic assessment of long-term silvicultural treatments within an eastern hemlock (Tsuga canadensis) forest, and computer-based simulated harvests of a genetically mapped eastern white pine (Pinus strobus) stand, we found that the selective removal of trees can alter gene frequencies. Due to an association with phenotypic characteristics used to guide harvests, the frequencies of rare alleles appeared particularly vulnerable to manipulation. Depending on the selection criteria used, rare allele frequencies either remained steady, decreased, or increased relative to study controls. Although harvest-associated genetic losses are possible, our data suggests that management can also sustain or enhance genetic richness. Similar to studies within temperate ecosystems, recent research in tropical forests underscores the potential influence of harvesting on the genetics of tree populations. In addition to efforts to reduce controllable sources of ecosystem stress (e.g., high pollutant exposures), management options should be evaluated that may bolster forest ecosystem resilience by preserving levels of genetic diversity within forests

Growth of canopy red oak near its northern range limit: current trends, potential drivers, and implications for the future

Red oak (Quercus rubra L.) is projected to expand into the northern hardwood forest over the coming century. We explored the connection between red oak basal area growth and a number of factors (tree age and size, stand dynamics, site elevation, and climate and acid deposition variables) for 213 trees in 11 plots throughout Vermont, USA. Red oak growth generally increased over the course of the chronology (1935–2014) and has been particularly high in recent decades. Growth differed among elevational groups but did not differ between age or size groups. Summer moisture metrics were consistently and positively associated with growth, whereas fall moisture was associated with reduced growth in recent decades. Higher summer temperatures were often negatively associated with growth, though there was evidence that low temperatures in the summer (higher elevations) and fall (lower elevations) constrain growth. Several pollution metrics were associated with reduced growth, a surprising result for a species not known to be sensitive to inputs of acid deposition that have predisposed other species in the region to decline. While red oak growth is currently robust, increases in summer temperatures, reductions in growing season precipitation, or increases in fall precipitation could reduce future growth potential.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author