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1.  Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients 
Annals of Botany  2013;112(3):623-631.
Background and Aims
The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.
Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).
Key Results
Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.
Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.
PMCID: PMC3718216  PMID: 23788748
Carbon supply; elevational gradient; Larix decidua; Nothofagus betuloides; Nothofagus pumilio; Patagonia; Pinus cembra; Pinus sylvestris; Swiss Alps; Alpine treeline
2.  Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees 
Annals of Botany  2011;108(7):1279-1286.
Background and Aims
Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.
Focus is given to relationships between tree height and top-shoot traits, which include the shoot's leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot's stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.
Key Results
Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.
In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.
PMCID: PMC3197456  PMID: 21914698
Allometry; current-year leader shoot; hierarchical Bayesian model; pipe model; tree height; water stress
3.  Responses of sap flow, leaf gas exchange and growth of hybrid aspen to elevated atmospheric humidity under field conditions 
AoB Plants  2014;6:plu021.
We demonstrate that higher air humidity mitigates the effect of low soil water availability on broadleaved trees during dry years by reducing stomatal limitation to photosynthesis, allowing higher net photosynthetic rates and supporting higher growth rates. At the same time, rising atmospheric humidity increases sensitivity of canopy conductance to water deficit and reduces the responsiveness of intrinsic water-use efficiency. The results imply that a future rise in atmospheric humidity at high latitudes may be disadvantageous in evenly rainy/humid years and expose trees to higher dehydration risk during weather extremes, although mitigating the impact of soil water deficit under moderate drought.
An increase in average air temperature and frequency of rain events is predicted for higher latitudes by the end of the 21st century, accompanied by a probable rise in air humidity. We currently lack knowledge on how forest trees acclimate to rising air humidity in temperate climates. We analysed the leaf gas exchange, sap flow and growth characteristics of hybrid aspen (Populus tremula × P. tremuloides) trees growing at ambient and artificially elevated air humidity in an experimental forest plantation situated in the hemiboreal vegetation zone. Humidification manipulation did not affect the photosynthetic capacity of plants, but did affect stomatal responses: trees growing at elevated air humidity had higher stomatal conductance at saturating photosynthetically active radiation (gs sat) and lower intrinsic water-use efficiency (IWUE). Reduced stomatal limitation of photosynthesis in trees grown at elevated air humidity allowed slightly higher net photosynthesis and relative current-year height increments than in trees at ambient air humidity. Tree responses suggest a mitigating effect of higher air humidity on trees under mild water stress. At the same time, trees at higher air humidity demonstrated a reduced sensitivity of IWUE to factors inducing stomatal closure and a steeper decline in canopy conductance in response to water deficit, implying higher dehydration risk. Despite the mitigating impact of increased air humidity under moderate drought, a future rise in atmospheric humidity at high latitudes may be disadvantageous for trees during weather extremes and represents a potential threat in hemiboreal forest ecosystems.
PMCID: PMC4052457  PMID: 24887000
Atmospheric humidity; canopy conductance; climate change; net photosynthesis; photosynthetic capacity; relative stomatal limitation; stomatal conductance; water-use efficiency.
4.  Stable oxygen isotope and flux partitioning demonstrates understory of an oak savanna contributes up to half of ecosystem carbon and water exchange 
Semi-arid ecosystems contribute about 40% to global net primary production (GPP) even though water is a major factor limiting carbon uptake. Evapotranspiration (ET) accounts for up to 95% of the water loss and in addition, vegetation can also mitigate drought effects by altering soil water distribution. Hence, partitioning of carbon and water fluxes between the soil and vegetation components is crucial to gain mechanistic understanding of vegetation effects on carbon and water cycling. However, the possible impact of herbaceous vegetation in savanna type ecosystems is often overlooked. Therefore, we aimed at quantifying understory vegetation effects on the water balance and productivity of a Mediterranean oak savanna. ET and net ecosystem CO2 exchange (NEE) were partitioned based on flux and stable oxygen isotope measurements and also rain infiltration was estimated. The understory vegetation contributed importantly to total ecosystem ET and GPP with a maximum of 43 and 51%, respectively. It reached water-use efficiencies (WUE; ratio of carbon gain by water loss) similar to cork-oak trees. The understory vegetation inhibited soil evaporation (E) and, although E was large during wet periods, it did not diminish WUE during water-limited times. The understory strongly increased soil water infiltration, specifically following major rain events. At the same time, the understory itself was vulnerable to drought, which led to an earlier senescence of the understory growing under trees as compared to open areas, due to competition for water. Thus, beneficial understory effects are dominant and contribute to the resilience of this ecosystem. At the same time the vulnerability of the understory to drought suggests that future climate change scenarios for the Mediterranean basin threaten understory development. This in turn will very likely diminish beneficial understory effects like infiltration and ground water recharge and therefore ecosystem resilience to drought.
PMCID: PMC4188126  PMID: 25339970
partitioning; stable oxygen isotopes; evapotranspiration; savanna; dry-land ecosystems; net ecosystem CO2 exchange; water-use efficiency; soil infiltration
5.  Millennium-Scale Crossdating and Inter-Annual Climate Sensitivities of Standing California Redwoods 
PLoS ONE  2014;9(7):e102545.
Extremely decay-resistant wood and fire-resistant bark allow California’s redwoods to accumulate millennia of annual growth rings that can be useful in biological research. Whereas tree rings of Sequoiadendron giganteum (SEGI) helped formalize the study of dendrochronology and the principle of crossdating, those of Sequoia sempervirens (SESE) have proven much more difficult to decipher, greatly limiting dendroclimatic and other investigations of this species. We overcame these problems by climbing standing trees and coring trunks at multiple heights in 14 old-growth forest locations across California. Overall, we sampled 1,466 series with 483,712 annual rings from 120 trees and were able to crossdate 83% of SESE compared to 99% of SEGI rings. Standard and residual tree-ring chronologies spanning up to 1,685 years for SESE and 1,538 years for SEGI were created for each location to evaluate crossdating and to examine correlations between annual growth and climate. We used monthly values of temperature, precipitation, and drought severity as well as summer cloudiness to quantify potential drivers of inter-annual growth variation over century-long time series at each location. SESE chronologies exhibited a latitudinal gradient of climate sensitivities, contrasting cooler northern rainforests and warmer, drier southern forests. Radial growth increased with decreasing summer cloudiness in northern rainforests and a central SESE location. The strongest dendroclimatic relationship occurred in our southernmost SESE location, where radial growth correlated negatively with dry summer conditions and exhibited responses to historic fires. SEGI chronologies showed negative correlations with June temperature and positive correlations with previous October precipitation. More work is needed to understand quantitative relationships between SEGI radial growth and moisture availability, particularly snowmelt. Tree-ring chronologies developed here for both redwood species have numerous scientific applications, including determination of tree ages, accurate dating of fire-return intervals, archaeology, analyses of stable isotopes, long-term climate reconstructions, and quantifying rates of carbon sequestration.
PMCID: PMC4102271  PMID: 25029026
6.  Pan-Tropical Analysis of Climate Effects on Seasonal Tree Growth 
PLoS ONE  2014;9(3):e92337.
Climate models predict a range of changes in tropical forest regions, including increased average temperatures, decreased total precipitation, reduced soil moisture and alterations in seasonal climate variations. These changes are directly related to the increase in anthropogenic greenhouse gas concentrations, primarily CO2. Assessing seasonal forest growth responses to climate is of utmost importance because woody tissues, produced by photosynthesis from atmospheric CO2, water and light, constitute the main component of carbon sequestration in the forest ecosystem. In this paper, we combine intra-annual tree growth measurements from published tree growth data and the corresponding monthly climate data for 25 pan-tropical forest sites. This meta-analysis is designed to find the shared climate drivers of tree growth and their relative importance across pan-tropical forests in order to improve carbon uptake models in a global change context. Tree growth reveals significant intra-annual seasonality at seasonally dry sites or in wet tropical forests. Of the overall variation in tree growth, 28.7% was explained by the site effect, i.e. the tree growth average per site. The best predictive model included four climate variables: precipitation, solar radiation (estimated with extrasolar radiation reaching the atmosphere), temperature amplitude and relative soil water content. This model explained more than 50% of the tree growth variations across tropical forests. Precipitation and solar radiation are the main seasonal drivers of tree growth, causing 19.8% and 16.3% of the tree growth variations. Both have a significant positive association with tree growth. These findings suggest that forest productivity due to tropical tree growth will be reduced in the future if climate extremes, such as droughts, become more frequent.
PMCID: PMC3966775  PMID: 24670981
7.  The Influence of Recent Climate Change on Tree Height Growth Differs with Species and Spatial Environment 
PLoS ONE  2011;6(2):e14691.
Tree growth has been reported to increase in response to recent global climate change in controlled and semi-controlled experiments, but few studies have reported response of tree growth to increased temperature and atmospheric carbon dioxide (CO2) concentration in natural environments. This study addresses how recent global climate change has affected height growth of trembling aspen (Populus tremuloides Michx) and black spruce (Picea mariana Mill B.S.) in their natural environments. We sampled 145 stands dominated by aspen and 82 dominated by spruce over the entire range of their distributions in British Columbia, Canada. These stands were established naturally after fire between the 19th and 20th centuries. Height growth was quantified as total heights of sampled dominant and co-dominant trees at breast-height age of 50 years. We assessed the relationships between 50-year height growth and environmental factors at both spatial and temporal scales. We also tested whether the tree growth associated with global climate change differed with spatial environment (latitude, longitude and elevation). As expected, height growth of both species was positively related to temperature variables at the regional scale and with soil moisture and nutrient availability at the local scale. While height growth of trembling aspen was not significantly related to any of the temporal variables we examined, that of black spruce increased significantly with stand establishment date, the anomaly of the average maximum summer temperature between May-August, and atmospheric CO2 concentration, but not with the Palmer Drought Severity Index. Furthermore, the increase of spruce height growth associated with recent climate change was higher in the western than in eastern part of British Columbia. This study demonstrates that the response of height growth to recent climate change, i.e., increasing temperature and atmospheric CO2 concentration, did not only differ with tree species, but also their growing spatial environment.
PMCID: PMC3040169  PMID: 21358817
8.  Contrasting trait syndromes in angiosperms and conifers are associated with different responses of tree growth to temperature on a large scale 
Recent large-scale studies of tree growth in the Iberian Peninsula reported contrasting positive and negative effects of temperature in Mediterranean angiosperms and conifers. Here we review the different hypotheses that may explain these trends and propose that the observed contrasting responses of tree growth to temperature in this region could be associated with a continuum of trait differences between angiosperms and conifers. Angiosperm and conifer trees differ in the effects of phenology in their productivity, in their growth allometry, and in their sensitivity to competition. Moreover, angiosperms and conifers significantly differ in hydraulic safety margins, sensitivity of stomatal conductance to vapor-pressure deficit (VPD), xylem recovery capacity or the rate of carbon transfer. These differences could be explained by key features of the xylem such as non-structural carbohydrate content (NSC), wood parenchymal fraction or wood capacitance. We suggest that the reviewed trait differences define two contrasting ecophysiological strategies that may determine qualitatively different growth responses to increased temperature and drought. Improved reciprocal common garden experiments along altitudinal or latitudinal gradients would be key to quantify the relative importance of the different hypotheses reviewed. Finally, we show that warming impacts in this area occur in an ecological context characterized by the advance of forest succession and increased dominance of angiosperm trees over extensive areas. In this context, we examined the empirical relationships between the responses of tree growth to temperature and hydraulic safety margins in angiosperm and coniferous trees. Our findings suggest a future scenario in Mediterranean forests characterized by contrasting demographic responses in conifer and angiosperm trees to both temperature and forest succession, with increased dominance of angiosperm trees, and particularly negative impacts in pines.
PMCID: PMC3797994  PMID: 24146668
conifers; angiosperms; functional traits; mediterranean ecosystems; drought; temperature; carbon metabolism; growth
9.  An Integrated Model of Environmental Effects on Growth, Carbohydrate Balance, and Mortality of Pinus ponderosa Forests in the Southern Rocky Mountains 
PLoS ONE  2013;8(11):e80286.
Climate-induced tree mortality is an increasing concern for forest managers around the world. We used a coupled hydrologic and ecosystem carbon cycling model to assess temperature and precipitation impacts on productivity and survival of ponderosa pine (Pinus ponderosa). Model predictions were evaluated using observations of productivity and survival for three ponderosa pine stands located across an 800 m elevation gradient in the southern Rocky Mountains, USA, during a 10-year period that ended in a severe drought and extensive tree mortality at the lowest elevation site. We demonstrate the utility of a relatively simple representation of declines in non-structural carbohydrate (NSC) as an approach for estimating patterns of ponderosa pine vulnerability to drought and the likelihood of survival along an elevation gradient. We assess the sensitivity of simulated net primary production, NSC storage dynamics, and mortality to site climate and soil characteristics as well as uncertainty in the allocation of carbon to the NSC pool. For a fairly wide set of assumptions, the model estimates captured elevational gradients and temporal patterns in growth and biomass. Model results that best predict mortality risk also yield productivity, leaf area, and biomass estimates that are qualitatively consistent with observations across the sites. Using this constrained set of parameters, we found that productivity and likelihood of survival were equally dependent on elevation-driven variation in temperature and precipitation. Our results demonstrate the potential for a coupled hydrology-ecosystem carbon cycling model that includes a simple model of NSC dynamics to predict drought-related mortality. Given that increases in temperature and in the frequency and severity of drought are predicted for a broad range of ponderosa pine and other western North America conifer forest habitats, the model potentially has broad utility for assessing ecosystem vulnerabilities.
PMCID: PMC3840024  PMID: 24282532
10.  Leaf-traits and growth allometry explain competition and differences in response to climatic change in a temperate forest landscape: a simulation study 
Annals of Botany  2011;108(5):885-894.
Background and Aims
The ability to simulate plant competition accurately is essential for plant functional type (PFT)-based models used in climate-change studies, yet gaps and uncertainties remain in our understanding of the details of the competition mechanisms and in ecosystem responses at a landscape level. This study examines secondary succession in a temperate deciduous forest in eastern China with the aim of determining if competition between tree types can be explained by differences in leaf ecophysiological traits and growth allometry, and whether ecophysiological traits and habitat spatial configurations among PFTs differentiate their responses to climate change.
A temperate deciduous broadleaved forest in eastern China was studied, containing two major vegetation types dominated by Quercus liaotungensis (OAK) and by birch/poplar (Betula platyphylla and Populus davidiana; BIP), respectively. The Terrestrial Ecosystem Simulator (TESim) suite of models was used to examine carbon and water dynamics using parameters measured at the site, and the model was evaluated against long-term data collected at the site.
Key Results
Simulations indicated that a higher assimilation rate for the BIP vegetation than OAK led to the former's dominance during early successional stages with relatively low competition. In middle/late succession with intensive competition for below-ground resources, BIP, with its lower drought tolerance/resistance and smaller allocation to leaves/roots, gave way to OAK. At landscape scale, predictions with increased temperature extrapolated from existing weather records resulted in increased average net primary productivity (NPP; +19 %), heterotrophic respiration (+23 %) and net ecosystem carbon balance (+17 %). The BIP vegetation in higher and cooler habitats showed 14 % greater sensitivity to increased temperature than the OAK at lower and warmer locations.
Drought tolerance/resistance and morphology-related allocation strategy (i.e. more allocation to leaves/roots) played key roles in the competition between the vegetation types. The overall site-average impacts of increased temperature on NPP and carbon stored in plants were found to be positive, despite negative effects of increased respiration and soil water stress, with such impacts being more significant for BIP located in higher and cooler habitats.
PMCID: PMC3177688  PMID: 21835816
Succession; watershed; ecophysiological trait; landscape carbon dynamics; temperate forests; competition; simulation; Quercus liaotungensis; Betula platyphylla; Populus davidiana
11.  Dynamics of non-structural carbohydrates in three Mediterranean woody species following long-term experimental drought 
Stored non-structural carbohydrates (NSC) have been proposed as a key determinant of drought resistance in plants. However, the evidence for this role is controversial, as it comes mostly from observational, short-term studies. Here, we take advantage of a long-term experimental throughfall reduction to elucidate the response of NSC to increased drought 14 years after the beginning of the treatment in three Mediterranean resprouter trees (Quercus ilex L., Arbutus unedo L. and Phillyrea latifolia L.). In addition, we selected 20 Q. ilex individuals outside the experimental plots to directly assess the relationship between defoliation and NSC at the individual level. We measured the seasonal course of NSC concentrations in leaves, branches and lignotuber in late winter, late spring, summer, and autumn 2012. Total concentrations of NSC were highest in the lignotuber for all species. In the long-term drought experiment we found significant depletion in concentrations of total NSC in treatment plots only in the lignotuber of A. unedo. At the same time, A. unedo was the only species showing a significant reduction in BAI under the drought treatment during the 14 years of the experiment. By contrast, Q. ilex just reduced stem growth only during the first 4 years of treatment and P. latifolia remained unaffected over the whole study period. However, we found a clear association between the concentrations of NSC and defoliation in Q. ilex individuals sampled outside the experimental plots, with lower total concentrations of NSC and lower proportion of starch in defoliated individuals. Taken together, our results suggest that stabilizing processes, probably at the stand level, may have been operating in the long-term to mitigate any impact of drought on NSC levels, and highlight the necessity to incorporate long-term experimental studies of plant responses to drought.
PMCID: PMC3795346  PMID: 24130568
drought; crown condition; growth; long-term; non-structural carbohydrates; starch; throughfall manipulation
12.  Expression of Multiple Resistance Genes Enhances Tolerance to Environmental Stressors in Transgenic Poplar (Populus × euramericana ‘Guariento’) 
PLoS ONE  2011;6(9):e24614.
Commercial and non-commercial plants face a variety of environmental stressors that often cannot be controlled. In this study, transgenic hybrid poplar (Populus × euramericana ‘Guariento’) harboring five effector genes (vgb, SacB, JERF36, BtCry3A and OC-I) were subjected to drought, salinity, waterlogging and insect stressors in greenhouse or laboratory conditions. Field trials were also conducted to investigate long-term effects of transgenic trees on insects and salt tolerance in the transformants. In greenhouse studies, two transgenic lines D5-20 and D5-21 showed improved growth, as evidenced by greater height and basal diameter increments and total biomass relative to the control plants after drought or salt stress treatments. The improved tolerance to drought and salt was primarily attributed to greater instantaneous water use efficiency (WUEi) in the transgenic trees. The chlorophyll concentrations tended to be higher in the transgenic lines under drought or saline conditions. Transformed trees in drought conditions accumulated more fructan and proline and had increased Fv/Fm ratios (maximum quantum yield of photosystem II) under waterlogging stress. Insect-feeding assays in the laboratory revealed a higher total mortality rate and lower exuviation index of leaf beetle [Plagiodera versicolora (Laicharting)] larvae fed with D5-21 leaves, suggesting enhanced insect resistance in the transgenic poplar. In field trials, the dominance of targeted insects on 2-year-old D5-21 transgenic trees was substantially lower than that of the controls, indicating enhanced resistance to Coleoptera. The average height and DBH (diameter at breast height) of 2.5-year-old transgenic trees growing in naturally saline soil were 3.80% and 4.12% greater than those of the control trees, but these increases were not significant. These results suggested that multiple stress-resistance properties in important crop tree species could be simultaneously improved, although additional research is needed to fully understand the relationships between the altered phenotypes and the function of each transgene in multigene transformants.
PMCID: PMC3170361  PMID: 21931776
13.  Spatial and seasonal variations in mobile carbohydrates in Pinus cembra in the timberline ecotone of the Central Austrian Alps 
European journal of forest research  2011;130(2):173-179.
To test whether the altitudinal limit of tree growth is determined by carbons shortage or by a limitation in growth we investigated non structural carbohydrates and their components starch and total soluble sugars in Pinus cembra trees along an elevational gradient in the timberline ecotone of the Central Austrian Alps. NSC contents in needles, branches, stems, and coarse roots were measured throughout an entire growing season. At the tissue level NSC contents were not significantly more abundant in treeline trees as compared to trees at lower elevations. Along our 425 m elevational transect from the closed forest to the treeline we failed to find a stable elevational trend in the total NSC pool of entire trees and observed within season increases in the tree’s NSC pool that can be attributed to an altitudinal increase in leaf mass as needles contained the largest NSC fraction of the whole tree NSC pool. Furthermore, whole tree NSC contents were positively correlated with net photosynthetic capacity. Although our observed NSC characteristics do not support the hypothesis that tree life at their upper elevational limit is determined by an insufficient carbon balance we found no consistent confirmation for the sink limitation hypothesis.
PMCID: PMC3191523  PMID: 22003357
Non structural carbohydrates; seasonal variation; elevational gradient; timberline ecotone; treeline formation; treelife limitation
14.  Variation of Maximum Tree Height and Annual Shoot Growth of Smith Fir at Various Elevations in the Sygera Mountains, Southeastern Tibetan Plateau 
PLoS ONE  2012;7(3):e31725.
Little is known about tree height and height growth (as annual shoot elongation of the apical part of vertical stems) of coniferous trees growing at various altitudes on the Tibetan Plateau, which provides a high-elevation natural platform for assessing tree growth performance in relation to future climate change. We here investigated the variation of maximum tree height and annual height increment of Smith fir (Abies georgei var. smithii) in seven forest plots (30 m×40 m) along two altitudinal transects between 3,800 m and 4,200/4,390 m above sea level (a.s.l.) in the Sygera Mountains, southeastern Tibetan Plateau. Four plots were located on north-facing slopes and three plots on southeast-facing slopes. At each site, annual shoot growth was obtained by measuring the distance between successive terminal bud scars along the main stem of 25 trees that were between 2 and 4 m high. Maximum/mean tree height and mean annual height increment of Smith fir decreased with increasing altitude up to the tree line, indicative of a stress gradient (the dominant temperature gradient) along the altitudinal transect. Above-average mean minimum summer (particularly July) temperatures affected height increment positively, whereas precipitation had no significant effect on shoot growth. The time series of annual height increments of Smith fir can be used for the reconstruction of past climate on the southeastern Tibetan Plateau. In addition, it can be expected that the rising summer temperatures observed in the recent past and anticipated for the future will enhance Smith fir's growth throughout its altitudinal distribution range.
PMCID: PMC3291607  PMID: 22396738
15.  No evidence for depletion of carbohydrate pools in Scots pine (Pinus sylvestris L.) under drought stress 
The physiological mechanisms leading to Scots pine (Pinus sylvestris L.) decline in the dry inner Alpine valleys are still unknown. Testing the carbon starvation hypothesis, we analysed the seasonal course of mobile carbohydrate pools (NSC) of Scots pine growing at a xeric and a dry-mesic site within an inner Alpine dry valley (750 m a.s.l., Tyrol, Austria) during the year 2009, which was characterized by exceptional soil dryness. Although, soil moisture content dropped to c. 10% at both sites during the growing season, NSC concentrations were rising in all tissues (branch, stem, root) till end of July, except in needles where maxima were reached around bud break. NSC concentrations were not significantly different in the analysed tissues at the xeric and the dry-mesic site. At the dry-mesic site NSC concentrations in the above ground tree biomass were significantly higher during the period of radial growth. An accumulation of NSC in roots at the end of July indicates a change in carbon allocation after an early cessation in above ground growth, possibly due to elevated below ground carbon demand. In conclusion our results revealed that extensive soil dryness during the growing season did not lead to carbon depletion. However, even though C-reserves were not exhausted, a sequestration of carbohydrate pools during drought periods might lead to deficits in carbon supply that weaken tree vigour and drive tree mortality.
PMCID: PMC3427021  PMID: 21974742
non-structural carbohydrates; Scots pine; drought; dry inner Alpine valley; carbon starvation; tree mortality
16.  Drought responses of flood-tolerant trees in Amazonian floodplains 
Annals of Botany  2009;105(1):129-139.
Flood-tolerant tree species of the Amazonian floodplain forests are subjected to an annual dry period of variable severity imposed when low river-water levels coincide with minimal precipitation. Although the responses of these species to flooding have been examined extensively, their responses to drought, in terms of phenology, growth and physiology, have been neglected hitherto, although some information is found in publications that focus on flooding.
The present review examines the dry phase of the annual flooding cycle. It consolidates existing knowledge regarding responses to drought among adult trees and seedlings of many Amazonian floodplain species.
Main Findings
Flood-tolerant species display variable physiological responses to dry periods and drought that indicate desiccation avoidance, such as reduced photosynthetic activity and reduced root respiration. However, tolerance and avoidance strategies for drought vary markedly among species. Drought can substantially decrease growth, biomass and photosynthetic activity among seedlings in field and laboratory studies. When compared with the responses to flooding, drought can impose higher seedling mortality and slower growth rates, especially among evergreen species. Results indicate that tolerance and avoidance strategies for drought vary markedly between species. Both seedling recruitment and photosynthetic activity are affected by drought,
For many species, the effects of drought can be as important as flooding for survival and growth, particularly at the seedling phase of establishment, ultimately influencing species composition. In the context of climate change and predicted decreases in precipitation in the Amazon Basin, the effects of drought on plant physiology and species distribution in tropical floodplain forest ecosystems should not be overlooked.
PMCID: PMC2794061  PMID: 19880423
Drought responses; Amazonia; floodplain forests; tree ecology; várzea
17.  Linking carbon supply to root cell-wall chemistry and mechanics at high altitudes in Abies georgei 
Annals of Botany  2010;107(2):311-320.
Background and Aims
The mobile carbon supply to different compartments of a tree is affected by climate, but its impact on cell-wall chemistry and mechanics remains unknown. To understand better the variability in root growth and biomechanics in mountain forests subjected to substrate mass movement, we investigated root chemical and mechanical properties of mature Abies georgei var. smithii (Smith fir) growing at different elevations on the Tibet–Qinghai Plateau.
Thin and fine roots (0·1–4·0 mm in diameter) were sampled at three different elevations (3480, 3900 and 4330 m, the last corresponding to the treeline). Tensile resistance of roots of different diameter classes was measured along with holocellulose and non-structural carbon (NSC) content.
Key Results
The mean force necessary to break roots in tension decreased significantly with increasing altitude and was attributed to a decrease in holocellulose content. Holocellulose was significantly lower in roots at the treeline (29·5 ± 1·3 %) compared with those at 3480 m (39·1 ± 1·0 %). Roots also differed significantly in NSC, with 35·6 ± 4·1 mg g−1 dry mass of mean total soluble sugars in roots at 3480 m and 18·8 ± 2·1 mg g−1 dry mass in roots at the treeline.
Root mechanical resistance, holocellulose and NSC content all decreased with increasing altitude. Holocellulose is made up principally of cellulose, the biosynthesis of which depends largely on NSC supply. Plants synthesize cellulose when conditions are optimal and NSC is not limiting. Thus, cellulose synthesis in the thin and fine roots measured in our study is probably not a priority in mature trees growing at very high altitudes, where climatic factors will be limiting for growth. Root NSC stocks at the treeline may be depleted through over-demand for carbon supply due to increased fine root production or winter root growth.
PMCID: PMC3025735  PMID: 21186240
Abies georgei; biomechanics; tensile resistance; cellulose content; non-structural carbon; root strength; Tibet–Qinghai; slope stability
18.  A Phenological Timetable of Oak Growth under Experimental Drought and Air Warming 
PLoS ONE  2014;9(2):e89724.
Climate change is expected to increase temperature and decrease summer precipitation in Central Europe. Little is known about how warming and drought will affect phenological patterns of oaks, which are considered to possess excellent adaptability to these climatic changes. Here, we investigated bud burst and intra-annual shoot growth of Quercus robur, Q. petraea and Q. pubescens grown on two different forest soils and exposed to air warming and drought. Phenological development was assessed over the course of three growing seasons. Warming advanced bud burst by 1–3 days °C−1 and led to an earlier start of intra-annual shoot growth. Despite this phenological shift, total time span of annual growth and shoot biomass were not affected. Drought changed the frequency and intensity of intra-annual shoot growth and advanced bud burst in the subsequent spring of a severe summer drought by 1–2 days. After re-wetting, shoot growth recovered within a few days, demonstrating the superior drought tolerance of this tree genus. Our findings show that phenological patterns of oaks are modified by warming and drought but also suggest that ontogenetic factors and/or limitations of water and nutrients counteract warming effects on the biomass and the entire span of annual shoot growth.
PMCID: PMC3933646  PMID: 24586988
19.  Growth-Mortality Relationships in Piñon Pine (Pinus edulis) during Severe Droughts of the Past Century: Shifting Processes in Space and Time 
PLoS ONE  2014;9(5):e92770.
The processes leading to drought-associated tree mortality are poorly understood, particularly long-term predisposing factors, memory effects, and variability in mortality processes and thresholds in space and time. We use tree rings from four sites to investigate Pinus edulis mortality during two drought periods in the southwestern USA. We draw on recent sampling and archived collections to (1) analyze P. edulis growth patterns and mortality during the 1950s and 2000s droughts; (2) determine the influence of climate and competition on growth in trees that died and survived; and (3) derive regression models of growth-mortality risk and evaluate their performance across space and time. Recent growth was 53% higher in surviving vs. dying trees, with some sites exhibiting decades-long growth divergences associated with previous drought. Differential growth response to climate partly explained growth differences between live and dead trees, with responses wet/cool conditions most influencing eventual tree status. Competition constrained tree growth, and reduced trees’ ability to respond to favorable climate. The best predictors in growth-mortality models included long-term (15–30 year) average growth rate combined with a metric of growth variability and the number of abrupt growth increases over 15 and 10 years, respectively. The most parsimonious models had high discriminatory power (ROC>0.84) and correctly classified ∼70% of trees, suggesting that aspects of tree growth, especially over decades, can be powerful predictors of widespread drought-associated die-off. However, model discrimination varied across sites and drought events. Weaker growth-mortality relationships and higher growth at lower survival probabilities for some sites during the 2000s event suggest a shift in mortality processes from longer-term growth-related constraints to shorter-term processes, such as rapid metabolic decline even in vigorous trees due to acute drought stress, and/or increases in the attack rate of both chronically stressed and more vigorous trees by bark beetles.
PMCID: PMC4008371  PMID: 24786646
20.  Alteration of Forest Structure Modifies the Distribution of Scale Insect, Stigmacoccus garmilleri, in Mexican Tropical Montane Cloud Forests 
Stigmacoccus garmilleri Foldi (Hemiptera: Margarodidae) is an ecologically important honeydew-producing scale insect associated with oak trees (Quercus spp.) in highland forests of Veracruz, Mexico. The honeydew exudates of S. garmilleri serve as a significant nutrient source to many species of birds, insects, and sooty molds. Oak trees found in the forest interior, forest edge, and those scattered in pasture areas support scale insect colonies, though the pattern of insect infestations on trees within these varying landscape types has not been elucidated. This study aims to describe the distribution of scale insect infestation and any distinctions in honeydew production based on tree location. Scale insect density, honeydew volume, and sugar concentration were surveyed throughout a continuous landscape that included both patches of forest and scattered pasture trees. In addition, the anal filament through which the honeydew drop is secreted was also measured and was experimentally removed to test and measure regrowth. Scale insect densities on tree trunks were greatest on pasture trees, while intermediate densities were found on trees at the forest edge, and low densities on interior forest trees, suggesting that trees in disturbed areas are more susceptible to scale insect infestation. Trees with small diameters at breast height had significantly higher insect densities than trees with medium to large diameters. Trunk aspect (North, South, East, and West) was not a significant determinant of scale insect density. In forested areas higher densities of scale insects were found at three meters height in comparison to lower heights. Sugar concentrations and drop volumes of honeydew in forest and pasture areas were not significantly different. However, scale-insect anal tubes/filaments were significantly longer in pasture than they were in forests. Sugar concentrations of honeydew appeared to be positively correlated with temperature and negatively correlated with relative humidity. Experiments indicated that anal filaments could grow approximately 4 mm every 24 hours, and average tube growth was significantly faster in pasture than in forest, suggesting that there may be a physiological effect on the insect due to landscape disturbance. The results obtained in this study describe the increases in scale insect infestation of trees with forest disturbance. The effect of these increased scale insect densities on the host tree physiology is still to be resolved.
PMCID: PMC3281392  PMID: 22239677
habitat fragmentation; land use change; Margarodidae; plant-animal interactions; Stigmacoccidae
21.  Struggle in the flood: tree responses to flooding stress in four tropical floodplain systems 
AoB Plants  2010;2010:plq003.
This review examines species diversity and structural, physiological and biochemical characteristics associated with survival of annual deep flooding of trees in four contrasting tropical floodplain ecosystems.
Background and aims
In the context of the 200th anniversary of Charles Darwin's birth in 1809, this study discusses the variation in structure and adaptation associated with survival and reproductive success in the face of environmental stresses in the trees of tropical floodplains.
We provide a comparative review on the responses to flooding stress in the trees of freshwater wetlands in tropical environments. The four large wetlands we evaluate are: (i) Central Amazonian floodplains in South America, (ii) the Okavango Delta in Africa, (iii) the Mekong floodplains of Asia and (iv) the floodplains of Northern Australia. They each have a predictable ‘flood pulse’. Although flooding height varies between the ecosystems, the annual pulse is a major driving force influencing all living organisms and a source of stress for which specialized adaptations for survival are required.
Main points
The need for trees to survive an annual flood pulse has given rise to a large variety of adaptations. However, phenological responses to the flood are similar in the four ecosystems. Deciduous and evergreen species respond with leaf shedding, although sap flow remains active for most of the year. Growth depends on adequate carbohydrate supply. Physiological adaptations (anaerobic metabolism, starch accumulation) are also required.
Data concerning the ecophysiology and adaptations of trees in floodplain forests worldwide are extremely scarce. For successful floodplain conservation, more information is needed, ideally through a globally co-ordinated study using reproducible comparative methods. In the light of climatic change, with increasing drought, decreased groundwater availability and flooding periodicities, this knowledge is needed ever more urgently to facilitate fast and appropriate management responses to large-scale environmental change.
PMCID: PMC2965040  PMID: 22476061
22.  Using plant growth modeling to analyze C source–sink relations under drought: inter- and intraspecific comparison 
The ability to assimilate C and allocate non-structural carbohydrates (NSCs) to the most appropriate organs is crucial to maximize plant ecological or agronomic performance. Such C source and sink activities are differentially affected by environmental constraints. Under drought, plant growth is generally more sink than source limited as organ expansion or appearance rate is earlier and stronger affected than C assimilation. This favors plant survival and recovery but not always agronomic performance as NSC are stored rather than used for growth due to a modified metabolism in source and sink leaves. Such interactions between plant C and water balance are complex and plant modeling can help analyzing their impact on plant phenotype. This paper addresses the impact of trade-offs between C sink and source activities and plant production under drought, combining experimental and modeling approaches. Two contrasted monocotyledonous species (rice, oil palm) were studied. Experimentally, the sink limitation of plant growth under moderate drought was confirmed as well as the modifications in NSC metabolism in source and sink organs. Under severe stress, when C source became limiting, plant NSC concentration decreased. Two plant models dedicated to oil palm and rice morphogenesis were used to perform a sensitivity analysis and further explore how to optimize C sink and source drought sensitivity to maximize plant growth. Modeling results highlighted that optimal drought sensitivity depends both on drought type and species and that modeling is a great opportunity to analyze such complex processes. Further modeling needs and more generally the challenge of using models to support complex trait breeding are discussed.
PMCID: PMC3817663  PMID: 24204372
drought; plant biomass accumulation; non-structural carbohydrate; source and sink regulation; functional structural plant model; rice; oil palm
23.  Tree-Ring Stable Isotopes Reveal Twentieth-Century Increases in Water-Use Efficiency of Fagus sylvatica and Nothofagus spp. in Italian and Chilean Mountains 
PLoS ONE  2014;9(11):e113136.
Changes in intrinsic water use efficiency (iWUE) were investigated in Fagus sylvatica and Nothofagus spp. over the last century. We combined dendrochronological methods with dual-isotope analysis to investigate whether atmospheric changes enhanced iWUE of Fagus and Nothofagus and tree growth (basal area increment, BAI) along latitudinal gradients in Italy and Chile. Post-maturation phases of the trees presented different patterns in δ13C, Δ13C, δ18O, Ci (internal CO2 concentration), iWUE, and BAI. A continuous enhancement in isotope-derived iWUE was observed throughout the twentieth century, which was common to all sites and related to changes in Ca (ambient CO2 concentration) and secondarily to increases in temperature. In contrast to other studies, we observed a general increasing trend of BAI, with the exception of F. sylvatica in Aspromonte. Both iWUE and BAI were uncoupled with the estimated drought index, which is in agreement with the absence of enduring decline in tree growth. In general, δ13C and δ18O showed a weak relationship, suggesting the major influence of photosynthetic rate on Ci and δ13C, and the minor contribution of the regulation of stomatal conductance to iWUE. The substantial warming observed during the twentieth century did not result in a clear pattern of increased drought stress along these latitudinal transects, because of the variability in temporal trends of precipitation and in specific responses of populations.
PMCID: PMC4232607  PMID: 25398040
24.  Recent Widespread Tree Growth Decline Despite Increasing Atmospheric CO2 
PLoS ONE  2010;5(7):e11543.
The synergetic effects of recent rising atmospheric CO2 and temperature are expected to favor tree growth in boreal and temperate forests. However, recent dendrochronological studies have shown site-specific unprecedented growth enhancements or declines. The question of whether either of these trends is caused by changes in the atmosphere remains unanswered because dendrochronology alone has not been able to clarify the physiological basis of such trends.
Methodology/Principal Findings
Here we combined standard dendrochronological methods with carbon isotopic analysis to investigate whether atmospheric changes enhanced water use efficiency (WUE) and growth of two deciduous and two coniferous tree species along a 9° latitudinal gradient across temperate and boreal forests in Ontario, Canada. Our results show that although trees have had around 53% increases in WUE over the past century, growth decline (measured as a decrease in basal area increment – BAI) has been the prevalent response in recent decades irrespective of species identity and latitude. Since the 1950s, tree BAI was predominantly negatively correlated with warmer climates and/or positively correlated with precipitation, suggesting warming induced water stress. However, where growth declines were not explained by climate, WUE and BAI were linearly and positively correlated, showing that declines are not always attributable to warming induced stress and additional stressors may exist.
Our results show an unexpected widespread tree growth decline in temperate and boreal forests due to warming induced stress but are also suggestive of additional stressors. Rising atmospheric CO2 levels during the past century resulted in consistent increases in water use efficiency, but this did not prevent growth decline. These findings challenge current predictions of increasing terrestrial carbon stocks under climate change scenarios.
PMCID: PMC2908114  PMID: 20657763
25.  How the timberline formed: altitudinal changes in stand structure and dynamics around the timberline in central Japan 
Annals of Botany  2012;109(6):1165-1174.
Background and Aims
Altitudinal timberlines are thought to move upward by global warming, a crucial topic in ecology. Tall tree species (the conifer Abies mariesii and the deciduous broad-leaved Betula ermanii) dominate the sub-alpine zone between 1600 and 2500 m a.s.l., the timberline, on Mount Norikura in central Japan. Dwarf pine Pinus pumila dominates above the timberline to near the summit (3026 m a.s.l.). This study evaluated how the timberline formed on Mount Norikura by examining altitudinal changes in stand structure and dynamics around the timberline.
One hundred and twenty-five plots of 10 m × 10 m were established around the timberline (2350–2600 m a.s.l.). Trunk diameter growth rate during 6 years was examined for A. mariesii, B. ermanii and P. pumila. Mortality during this period and mechanical damage scars on the trunks and branches due to strong wind and snow were examined for A. mariesii only.
Key Results
The density, maximum trunk height and diameter of A. mariesii in plots decreased with altitude. The maximum trunk height of B. ermanii decreased with altitude, but density and maximum trunk diameter did not decrease. In contrast, the density of P. pumila abruptly increased from around the timberline. A strong negative correlation was found between the densities of P. pumila and tall tree species, indicating their interspecific competition. Trunk diameter growth rates of A. mariesii and B. ermanii did not decrease with altitude, suggesting that these two tall tree species can grow at the timberline. The ratio of trees with mechanical damage scars increased with altitude for A. mariesii, a tendency more conspicuous for larger trees. The mortality of larger A. mariesii was also greater at higher altitude. Tall tree species may not increase their trunk height and survive around the timberline because of mechanical damage.
This study suggests that the altitudinal location of the timberline is mainly affected by mechanical damage due to strong wind and snow rather than by growth limitation due to low temperature. Therefore, the timberline would not move upward even under global warming if these growth and mortality characteristics do not change for a long time.
PMCID: PMC3336954  PMID: 22451598
Abies mariesii; Betula ermanii; climate change; disturbance; global warming; Pinus pumila; timberline

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