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1.  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
2.  Long-term changes in tree-ring – climate relationships at Mt. Patscherkofel (Tyrol, Austria) since the mid 1980s 
Trees (Berlin, Germany : West)  2008;22(1):31-40.
Although growth limitation of trees at Alpine and high-latitude timberlines by prevailing summer temperature is well established, loss of thermal response of radial tree growth during last decades has repeatedly been addressed. We examined long-term variability of climate-growth relationships in ring width chronologies of Stone pine (Pinus cembra L.) by means of moving response functions (MRF). The study area is situated in the timberline ecotone (c. 2000 – 2200 m a.s.l.) on Mt. Patscherkofel (Tyrol, Austria). Five site chronologies were developed within the ecotone with constant sample depth (≥ 19 trees) throughout most of the time period analysed. MRF calculated for the period 1866-1999 and 1901-1999 for c. 200 and c. 100 yr old stands, respectively, revealed that mean July temperature is the major and long-term stable driving force of Pinus cembra radial growth within the timberline ecotone. However, since the mid 1980s, radial growth in timberline and tree line chronologies strikingly diverges from the July temperature trend. This is probably a result of extreme climate events (e.g. low winter precipitation, late frost) and/or increasing drought stress on cambial activity. The latter assumption is supported by a < 10 % increase in annual increments of c. 50 yr old trees at the timberline and at the tree line in 2003 compared to 2002, when extraordinary hot and dry conditions prevailed during summer. Furthermore, especially during the second half of the 20th century, influence of climate variables on radial growth show abrupt fluctuations, which might also be a consequence of climate warming on tree physiology.
PMCID: PMC3083837  PMID: 21532976
Climate warming; moving response function; Pinus cembra; temperature sensitivity; tree ring
3.  Cambial activity and xylem cell development in Pinus cembra and Pinus sylvestris at their climatic limits in the Eastern Alps in 2007 
Phyton; annales rei botanicae  2011;51(2):299-313.
It has been frequently stressed that at distributional boundaries, like at the Alpine timberline and within dry inner Alpine environments, tree growth will be affected first by changing climate conditions. Climate in 2007 was characterized by the occurrence of exceptionally mild temperatures in spring (3.4 and 2.7 °C above long-term mean (LTM) at timberline and the valley sites, respectively) with an almost continuous drought period recorded in April and slightly warmer than average temperatures throughout summer (1.3 °C above LTM at both sites).
We compared temporal dynamics of cambial activity and xylem cell development in Pinus cembra at the Alpine timberline (1950 m a.s.l.) and Pinus sylvestris at a xeric inner Alpine site (750 m a.s.l.) by repeated cellular analyses of micro-cores (n = 5 trees/site). While onset of wood formation in P. sylvestris and P. cembra differed by about two weeks (12 and 27 April, respectively), maximum daily growth rates peaked on 6 May at the valley site and on 23 June at timberline. At both sites maximum tracheid production was reached prior to occurrence of more favourable climatic conditions during summer, i.e. an increase in precipitation and temperature. Xylem formation ended on 31 August and 28 October at the xeric site and at timberline, respectively.
This study demonstrates the plasticity of tree-ring formation along an altitudinal transect in response to water availability and temperature. Whether early achievement of maximum growth rates is an adaptation to cope with extreme environmental conditions prevailing at limits of tree growth needs to be analysed more closely by taking belowground carbon allocation into account.
PMCID: PMC3837289  PMID: 24273354
Alpine timberline; cambium; dry inner Alpine valley; intra-annual growth; Scots pine; Stone pine; wood anatomy; xylogenesis
4.  Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables 
Trees (Berlin, Germany : West)  2009;23(3):623-635.
We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1950 m a.s.l.), treeline (2110 m a.s.l.) and within the krummholz belt (2180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined.
At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5 °C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth.
The rate of xylem cell production determined in two weekly intervals during June through August 2006-2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline.
Results of this study revealed that spatial and temporal (i.e. year-to-year) variability in timing and dynamic of wood formation of Pinus cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.
PMCID: PMC3078619  PMID: 21509148
Cambium; intra-annual growth; Pinus cembra; temperature; tracheid production
5.  Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone 
Annals of forest science  2009;66(5):503.
Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.
PMCID: PMC3059571  PMID: 21423861
dendrometer; Pinus cembra; radial increment; treeline ecotone; xylem formation
6.  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
7.  Intra-annual dynamics of stem CO2 efflux in relation to cambial activity and xylem development in Pinus cembra 
Tree physiology  2009;29(5):641-649.
The relationship between stem CO2 efflux (ES), cambial activity and xylem production in Pinus cembra was determined at the timberline (1950 m a.s.l.) of the Central Austrian Alps, throughout one year. ES was measured continuously from June 2006 to August 2007 using an infrared gas-analysis system. Cambial activity and xylem production was determined by repeated microcore sampling of the developing tree ring and radial increment was monitored using automated point dendrometers. Aside of temperature, the number of living tracheids and cambial cells was predominantly responsible for ES: ES normalized to 10°C (ES10) was significantly correlated to number of living cells throughout the year (r2 = 0,574; p < 0,001). However, elevated ES and missing correlation between ES10 and xylem production was detected during cambial reactivation in April and during transition from active phase to rest, which occurred in August and lasted until early September. Results of this study indicate that (i) during seasonal variations in cambial activity non-linearity between ES and xylem production occurs and (ii) elevated metabolic activity during transition stages in the cambial activity-dormancy cycle influence the carbon budget of Pinus cembra. Daily radial stem increment was primarily influenced by the number of enlarging cells and was not correlated to ES.
PMCID: PMC3013296  PMID: 19203979
cambial reactivation; dormancy; Pinus cembra; radial stem growth; sap flow; stem CO2 efflux; stem respiration; xylem production
8.  Effects of atmospheric and climate change at the timberline of the Central European Alps 
Annals of forest science  2009;66(4):402.
This review considers potential effects of atmospheric change and climate warming within the timberline ecotone of the Central European Alps. After focusing on the impacts of ozone (O3) and rising atmospheric CO2 concentration, effects of climate warming on the carbon and water balance of timberline trees and forests will be outlined towards conclusions about changes in tree growth and treeline dynamics.Presently, ambient ground-level O3 concentrations do not exert crucial stress on adult conifers at the timberline of the Central European Alps. In response to elevated atmospheric CO2 Larix decidua showed growth increase, whereas no such response was found in Pinus uncinata. Overall climate warming appears as the factor responsible for the observed growth stimulation of timberline trees.Increased seedling re-establishment in the Central European Alps however, resulted from invasion into potential habitats rather than upward migration due to climate change, although seedlings will only reach tree size upon successful coupling with the atmosphere and thus loosing the beneficial microclimate of low stature vegetation.In conclusion, future climate extremes are more likely than the gradual temperature increase to control treeline dynamics in the Central European Alps.
PMCID: PMC3047780  PMID: 21379395
Alpine timberline; treeline; global warming; CO2; ozone; water balance; carbon gain
9.  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
10.  Field and controlled environment measurements show strong seasonal acclimation in photosynthesis and respiration potential in boreal Scots pine 
Understanding the seasonality of photosynthesis in boreal evergreen trees and its control by the environment requires separation of the instantaneous and slow responses, as well as the dynamics of light reactions, carbon reactions, and respiration. We determined the seasonality of photosynthetic light response and respiration parameters of Scots pine (Pinus sylvestris L.) in the field in southern Finland and in controlled laboratory conditions. CO2 exchange and chlorophyll fluorescence were measured in the field using a continuously operated automated chamber setup and fluorescence monitoring systems. We also carried out monthly measurements of photosynthetic light, CO2 and temperature responses in standard conditions with a portable IRGA and fluorometer instrument. The field and response measurements indicated strong seasonal variability in the state of the photosynthetic machinery with a deep downregulation during winter. Despite the downregulation, the photosynthetic machinery retained a significant capacity during winter, which was not visible in the field measurements. Light-saturated photosynthesis (Psat) and the initial slope of the photosynthetic light response (α) obtained in standard conditions were up to 20% of their respective summertime values. Respiration also showed seasonal acclimation with peak values of respiration in standard temperature in spring and decline in autumn. Spring recovery of all photosynthetic parameters could be predicted with temperature history. On the other hand, the operating quantum yield of photosystem II and the initial slope of photosynthetic light response stayed almost at the summertime level until late autumn while at the same time Psat decreased following the prevailing temperature. Comparison of photosynthetic parameters with the environmental drivers suggests that light and minimum temperature are also decisive factors in the seasonal acclimation of photosynthesis in boreal evergreen trees.
PMCID: PMC4264410  PMID: 25566291
gas exchange; chlorophyll fluorescence; annual cycle; Pmax
11.  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.
12.  Intra-annual variability of anatomical structure and δ13C values within tree rings of spruce and pine in alpine, temperate and boreal Europe 
Oecologia  2009;161(4):729-745.
Tree-ring width, wood density, anatomical structure and 13C/12C ratios expressed as δ13C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ13C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ13C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ13C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ13C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ13C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ13C and tree-ring width and climate are multi-factorial in seasonal climates.
PMCID: PMC2744769  PMID: 19653008
Dendrochonology; Carbohydrate storage; Picea abies; Pinus cembra; Pinus sylvestris; Tracheid lumen area; Wood density
13.  Drought sensitivity of three co-occurring conifers within a dry inner Alpine environment 
Trees (Berlin, Germany : West)  2013;27(1):61-69.
We applied dendroclimatological techniques to determine long-term stationarity of climate-growth relationships and recent growth trends of three widespread coniferous tree species of the central Austrian Alps, which grow intermixed at dry-mesic sites within a dry inner Alpine environment (750 m asl). Time series of annual increments were developed from > 120 mature trees of Picea abies, Larix decidua and Pinus sylvestris. Calculation of response functions for the period 1911 – 2009 revealed significant differences among species in response to climate variables. While precipitation in May – June favoured radial growth of Picea abies and Larix decidua, Pinus sylvestris growth mainly depended on April – May precipitation. P. abies growth was most sensitive to May – June temperature (inverse relationship). Moving response function coefficients indicated increasing drought sensitivity of all species in recent decades, which is related to a decline in soil moisture availability due to increasing stand density and tree size and higher evapotranspiration rates in a warmer climate. While recent trend in basal area increment (BAI) of L. decidua distinctly declined implying high vulnerability to drought stress, moderately shade-tolerant P. abies showed steadily increasing BAI and quite constant BAI was maintained in drought adapted P. sylvestris, although at lowest level of all species. We conclude that synergistic effects of stand dynamics and climate warming increased drought sensitivity, which changed competitive strength of co-occurring conifers due to differences in inherent adaptive capacity.
PMCID: PMC3750198  PMID: 23976821
Basal area increment; Dendroclimatology; Inner Alpine valley; Radial growth; Moving response function; Tree-ring analysis
14.  Holocene temperature variations at a high-altitude site in the Eastern Alps: a chironomid record from Schwarzsee ob Sölden, Austria 
Quaternary Science Reviews  2011;30(1-2):176-191.
Few well-dated, quantitative Holocene temperature reconstructions exist from high-altitude sites in the Central Eastern Alps. Here, we present a chironomid-based quantitative reconstruction of mean July air temperatures (TJuly) throughout the Holocene for a remote high-mountain lake, Schwarzsee ob Sölden, situated above the treeline at 2796 m a.s.l. in the Austrian Alps. Applying a chironomid-temperature inference model developed from lakes of the Alpine region to a high-resolution chironomid record from the lake provides evidence for early Holocene (ca 10000–8600 cal yr BP) TJuly of up to 8.5 °C, i.e. >4 °C above the modern (1977–2006) mean July temperature. The reconstruction reveals the so-called ‘8.2-ka cold event’ centered at ca 8250–8000 cal yr BP with temperatures ca 3 °C below the early-Holocene thermal maximum. Rather warm (ca 6 °C) and productive conditions prevailed during ca 7900–4500 cal yr BP. The chironomid record suggests a climate transition between ca 5200 and 4500 cal yr BP to cooler TJuly. A distinct cooling trend is evident from ca 4500 until ca 2500 cal yr BP. Thereafter, the study site experienced its coldest conditions (around 4 °C or less) throughout the rest of the Holocene, with the exception of the warming trend during the late 20th century. Beside other factors, the Northern Hemisphere summer insolation seems to be the major driving force for the long-term trends in TJuly at high altitudes in the Eastern Alps. Due to the extreme location of the lake and the limited temperature range represented by the applied calibration data set, the chironomid-based temperature reconstruction fails to track phases of the late-Holocene climatic history with TJuly cooler than 4 °C. Further chironomid-based palaeoclimate model and down-core studies are required to address this problem, provide more realistic TJuly estimates from undisturbed high-altitude lakes in the Alps, and extract a reliable regional temperature signal.
PMCID: PMC3021123  PMID: 21317974
15.  Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum. 
The aims of this paper are to review previously published palaeovegetation and independent palaeoclimatic datasets together with new results we present from dynamic vegetation model simulations and modern pollen rain studies to: (i) determine the responses of Amazonian ecosystems to changes in temperature, precipitation and atmospheric CO2 concentrations that occurred since the last glacial maximum (LGM), ca. 21 000 years ago; and (ii) use this long-term perspective to predict the likely vegetation responses to future climate change. Amazonia remained predominantly forested at the LGM, although the combination of reduced temperatures, precipitation and atmospheric CO2 concentrations resulted in forests structurally and floristically quite different from those of today. Cold-adapted Andean taxa mixed with rainforest taxa in central areas, while dry forest species and lianas probably became important in the more seasonal southern Amazon forests and savannahs expanded at forest-savannah ecotones. Net primary productivity (NPP) and canopy density were significantly lower than today. Evergreen rainforest distribution and NPP increased during the glacial-Holocene transition owing to ameliorating climatic and CO2 conditions. However, reduced precipitation in the Early-Mid-Holocene (ca. 8000-3600 years ago) caused widespread, frequent fires in seasonal southern Amazonia, causing increased abundance of drought-tolerant dry forest taxa and savannahs in ecotonal areas. Rainforests expanded once more in the Late Holocene owing to increased precipitation caused by greater austral summer insolation, although some of this forest expansion (e.g. in parts of the Bolivian Beni) is clearly caused by palaeo Indian landscape modification. The plant communities that existed during the Early-Mid-Holocene may provide insights into the kinds of vegetation response expected from similar increases in temperature and aridity predicted for the twenty-first century. We infer that ecotonal areas near the margins of the Amazon Basin are liable to be most sensitive to future environmental change and should therefore be targeted with conservation strategies that allow 'natural' species movements and plant community re-assortments to occur.
PMCID: PMC1693334  PMID: 15212099
16.  Frost resistance in alpine woody plants 
This report provides a brief review of key findings related to frost resistance in alpine woody plant species, summarizes data on their frost resistance, highlights the importance of freeze avoidance mechanisms, and indicates areas of future research. Freezing temperatures are possible throughout the whole growing period in the alpine life zone. Frost severity, comprised of both intensity and duration, becomes greater with increasing elevation and, there is also a greater probability, that small statured woody plants, may be insulated by snow cover. Several frost survival mechanisms have evolved in woody alpine plants in response to these environmental conditions. Examples of tolerance to extracellular freezing and freeze dehydration, life cycles that allow species to escape frost, and freeze avoidance mechanisms can all be found. Despite their specific adaption to the alpine environment, frost damage can occur in spring, while all alpine woody plants have a low risk of frost damage in winter. Experimental evidence indicates that premature deacclimation in Pinus cembra in the spring, and a limited ability of many species of alpine woody shrubs to rapidly reacclimate when they lose snow cover, resulting in reduced levels of frost resistance in the spring, may be particularly critical under the projected changes in climate. In this review, frost resistance and specific frost survival mechanisms of different organs (leaves, stems, vegetative and reproductive over-wintering buds, flowers, and fruits) and tissues are compared. The seasonal dynamics of frost resistance of leaves of trees, as opposed to woody shrubs, is also discussed. The ability of some tissues and organs to avoid freezing by supercooling, as visualized by high resolution infrared thermography, are also provided. Collectively, the report provides a review of the complex and diverse ways that woody plants survive in the frost dominated environment of the alpine life zone.
PMCID: PMC4249714  PMID: 25520725
ice susceptibility; freezing avoidance; frost dehardening; frost hardening; supercooling
17.  One-year experiment on the physiological response of the Mediterranean crustose coralline alga, Lithophyllum cabiochae, to elevated pCO2 and temperature 
Ecology and Evolution  2013;3(3):676-693.
The response of respiration, photosynthesis, and calcification to elevated pCO2 and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near-ambient conditions of irradiance, at ambient or elevated temperature (+3°C), and at ambient (ca. 400 μatm) or elevated pCO2 (ca. 700 μatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO2 but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO2. Conversely, photosynthesis was strongly affected by pCO2 with a decline under elevated pCO2 in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO2. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO2 but decreased with rising temperature under elevated pCO2. Surprisingly, the highest rates in summer were found under elevated pCO2 and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO2. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO2. However, there is likely a trade-off with other physiological processes. For example, photosynthesis declines in response to increased pCO2 under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.
PMCID: PMC3605855  PMID: 23533024
Calcification; coralligenous habitat; coralline algae; global warming; irradiance; ocean acidification; pCO2; photosynthesis; respiration; temperature
18.  A Preliminary Study of the Heating Effect of the Tibetan Plateau 
PLoS ONE  2013;8(7):e68750.
The immense and towering Tibetan Plateau acts as a heating source and, thus, deeply shapes the climate of the Eurasian continent and even the whole world. However, due to the scarcity of meteorological observation stations and very limited climatic data, little is quantitatively known about the heating effect of the plateau and its implications. This paper firstly collects climate data (2001–2007) from 109 observation stations and MODIS-based estimated monthly mean temperature data in the plateau and the neighboring Sichuan Basin, and conducts correlation and simple linear regression to reveal the altitudinal pattern of temperature. Then, according to the linear relationships of temperature and altitude for each month, it compares air temperature differences on the same elevation between the main plateau and surrounding mountains and the Sichuan Basin so as to quantify the heating effect and discuss its implication on timberline of the plateau. The results show that: 1) the heating effect of the plateau is significant. The temperature of the main plateau area was higher than that of free air on the same elevation above the neighboring areas; on the elevation of 4500 m (the main plateau), temperature is 1–6°C higher in the main Plateau than over the Sichuan Basin for different months and 5.9–10.7°C higher than in the Qilian Mountains in the northeastern corner of the plateau. 2) Even at altitudes of 5000–6000 m in the main Plateau, there are 4 months with a mean temperature above 0°C. The mean temperature of the warmest month (July) can reach 10°C at about 4600–4700 m. This may help explain why the highest timberline in the northern hemisphere is on the southeastern Tibetan Plateau.
PMCID: PMC3729943  PMID: 23935886
19.  Geographical and climatic limits of needle types of one- and two-needled pinyon pines 
Journal of Biogeography  2008;35(2):257-269.
The geographical extent and climatic tolerances of one- and two-needled pinyon pines (Pinus subsect. Cembroides) are the focus of questions in taxonomy, palaeoclimatology and modelling of future distributions. The identification of these pines, traditionally classified by one- versus two-needled fascicles, is complicated by populations with both one- and two-needled fascicles on the same tree, and the description of two more recently described one-needled varieties: the fallax-type and californiarum-type. Because previous studies have suggested correlations between needle anatomy and climate, including anatomical plasticity reflecting annual precipitation, we approached this study at the level of the anatomy of individual pine needles rather than species.
Western North America.
We synthesized available and new data from field and herbarium collections of needles to compile maps of their current distributions across western North America. Annual frequencies of needle types were compared with local precipitation histories for some stands. Historical North American climates were modelled on a c. 1-km grid using monthly temperature and precipitation values. A geospatial model (ClimLim), which analyses the effect of climate-modulated physiological and ecosystem processes, was used to rank the importance of seasonal climate variables in limiting the distributions of anatomical needle types.
The pinyon needles were classified into four distinct types based upon the number of needles per fascicle, needle thickness and the number of stomatal rows and resin canals. The individual needles fit well into four categories of needle types, whereas some trees exhibit a mixture of two needle types. Trees from central Arizona containing a mixture of Pinus edulis and fallax-type needles increased their percentage of fallax-type needles following dry years. All four needle types occupy broader geographical regions with distinctive precipitation regimes. Pinus monophylla and californiarum-type needles occur in regions with high winter precipitation. Pinus edulis and fallax-type needles are found in regions with high monsoon precipitation. Areas supporting californiarum-type and fallax-type needle distributions are additionally characterized by a more extreme May–June drought.
Main conclusions
These pinyon needle types seem to reflect the amount and seasonality of precipitation. The single needle fascicle characterizing the fallax type may be an adaptation to early summer or periodic drought, while the single needle of Pinus monophylla may be an adaptation to summer–autumn drought. Although the needles fit into four distinct categories, the parent trees are sometimes less easily classified, especially near their ancestral Pleistocene ranges in the Mojave and northern Sonoran deserts. The abundance of trees with both one- and two-needled fascicles in the zones between P. monophylla, P. edulis and fallax-type populations suggest that needle fascicle number is an unreliable characteristic for species classification. Disregarding needle fascicle number, the fallax-type needles are nearly identical to P. edulis, supporting Little’s (1968) initial classification of these trees as P. edulis var. fallax, while the californiarum-type needles have a distinctive morphology supporting Bailey’s (1987) classification of this tree as Pinus californiarum.
PMCID: PMC3001037  PMID: 21188300
Climate modelling; needle anatomy; Pinus edulis; Pinus monophylla; pinyon pines; species climate window; western North America
20.  The water balance of grassland ecosystems in the Austrian Alps 
The altitudinal variation of precipitation, evapotranspiration and runoff was quantified at 16 different grassland sites between 580 and 2550 m a.s.l. in the Austrian Alps. Along this altitudinal transect annual evapotranspiration decreased from roughly 690 mm at low elevation sites to 210 – 220 mm at the upper limit of the alpine grassland belt. A detailed analysis of the data showed that the observed reduction in the annual sum of evapotranspiration could be mainly explained by the altitudinal decline of the length of the snow free period (i.e. the vegetation period). Daily mean sums of evapotranspiration showed no altitudinal trend and averaged 2.2 mm d−1 independent of elevation, although the leaf area index, growing season mean air temperature, and vapour pressure deficit declined with increasing altitude. As precipitation increased with elevation, evapotranspiration seems to be of secondary importance when compared to runoff. Inter-annual variability of evapotranspiration was fairly low across contrasting dry and wet years (coefficient of variation = 7 %), indicating that even during dry years water availability was not limiting evapotranspiration.
PMCID: PMC3858980  PMID: 24348098
21.  Fluorescence measurements show stronger cold inhibition of photosynthetic light reactions in Scots pine compared to Norway spruce as well as during spring compared to autumn 
We studied the photosynthetic activity of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies [L.] Karst) in relation to air temperature changes from March 2013 to February 2014. We measured the chlorophyll fluorescence of approximately 50 trees of each species growing in southern Finland. Fluorescence was measured 1–3 times per week. We began by measuring shoots present in late winter (i.e., March 2013) before including new shoots once they started to elongate in spring. By July, when the spring shoots had achieved similar fluorescence levels to the older ones, we proceeded to measure the new shoots only. We analyzed the data by fitting a sigmoidal model containing four parameters to link sliding averages of temperature and fluorescence. A parameter defining the temperature range over which predicted fluorescence increased most rapidly was the most informative with in describing temperature dependence of fluorescence. The model generated similar fluorescence patterns for both species, but differences were observed for critical temperature and needle age. Down regulation of the light reaction was stronger in spring than in autumn. Pine showed more conservative control of the photosynthetic light reactions, which were activated later in spring and more readily attenuated in autumn. Under the assumption of a close correlation of fluorescence and photosynthesis, spruce should therefore benefit more than pine from the increased photosynthetic potential during warmer springs, but be more likely to suffer frost damage with a sudden cooling following a warm period. The winter of 2013–2014 was unusually mild and similar to future conditions predicted by global climate models. During the mild winter, the activity of photosynthetic light reactions of both conifers, especially spruce, remained high. Because light levels during winter are too low for photosynthesis, this activity may translate to a net carbon loss due to respiration.
PMCID: PMC4055857  PMID: 24982664
Scots pine; Norway spruce; photosynthesis; phenology; cold inhibition; climate change; frost tolerance
22.  Seasonal Variation in Respiration of 1‐year‐old Shoots of Scots Pine Exposed to Elevated Carbon Dioxide and Temperature for 4 Years 
Annals of Botany  2003;92(1):89-96.
Sixteen 20‐year‐old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed for 4 years in environment‐controlled chambers that maintained: (1) ambient conditions (CON); (2) elevated atmospheric CO2 concentration (ambient + 350 µmol mol–1; EC); (3) elevated temperature (ambient +2–6 °C; ET); or (4) elevated CO2 and elevated temperature (ECT). The dark respiration rates of 1‐year‐old shoots, from which needles had been partly removed, were measured over the growing season in the fourth year. In all treatments, the temperature coefficient of respiration, Q10, changed with season, being smaller during the growing season than at other times. Respiration rate varied diurnally and seasonally with temperature, being highest around mid‐summer and declining gradually thereafter. When measurements were made at the temperature of the chamber, respiration rates were reduced by the EC treatment relative to CON, but were increased by ET and ECT treatments. However, respiration rates at a reference temperature of 15 °C were reduced by ET and ECT treatments, reflecting a decreased capacity for respiration at warmer temperatures (negative acclimation). The interaction between season and treatment was not significant. Growth respiration did not differ between treatments, but maintenance respiration did, and the differences in mean daily respiration rate between the treatments were attributable to the maintenance component. We conclude that maintenance respiration should be considered when modelling respiratory responses to elevated CO2 and elevated temperature, and that increased atmospheric temperature is more important than increasing CO2 when assessing the carbon budget of pine forests under conditions of climate change.
PMCID: PMC4243642  PMID: 12763759
Scots pine; Pinus sylvestris L.; elevated CO2; elevated temperature; shoot respiration
23.  Composition and Structure of Pinus koraiensis Mixed Forest Respond to Spatial Climatic Changes 
PLoS ONE  2014;9(5):e97192.
Although some studies have indicated that climate changes can affect Pinus koraiensis mixed forest, the responses of composition and structure of Pinus koraiensis mixed forests to climatic changes are unknown and the key climatic factors controlling the composition and structure of Pinus koraiensis mixed forest are uncertain.
Methodology/principal findings
Field survey was conducted in the natural Pinus koraiensis mixed forests along a latitudinal gradient and an elevational gradient in Northeast China. In order to build the mathematical models for simulating the relationships of compositional and structural attributes of the Pinus koraiensis mixed forest with climatic and non-climatic factors, stepwise linear regression analyses were performed, incorporating 14 dependent variables and the linear and quadratic components of 9 factors. All the selected new models were computed under the +2°C and +10% precipitation and +4°C and +10% precipitation scenarios. The Max Temperature of Warmest Month, Mean Temperature of Warmest Quarter and Precipitation of Wettest Month were observed to be key climatic factors controlling the stand densities and total basal areas of Pinus koraiensis mixed forest. Increased summer temperatures and precipitations strongly enhanced the stand densities and total basal areas of broadleaf trees but had little effect on Pinus koraiensis under the +2°C and +10% precipitation scenario and +4°C and +10% precipitation scenario.
These results show that the Max Temperature of Warmest Month, Mean Temperature of Warmest Quarter and Precipitation of Wettest Month are key climatic factors which shape the composition and structure of Pinus koraiensis mixed forest. Although the Pinus koraiensis would persist, the current forests dominated by Pinus koraiensis in the region would all shift and become broadleaf-dominated forests due to the dramatic increase of broadleaf trees under the future global warming and increased precipitation.
PMCID: PMC4014611  PMID: 24810605
24.  Toward a Mechanistic Modeling of Nitrogen Limitation on Vegetation Dynamics 
PLoS ONE  2012;7(5):e37914.
Nitrogen is a dominant regulator of vegetation dynamics, net primary production, and terrestrial carbon cycles; however, most ecosystem models use a rather simplistic relationship between leaf nitrogen content and photosynthetic capacity. Such an approach does not consider how patterns of nitrogen allocation may change with differences in light intensity, growing-season temperature and CO2 concentration. To account for this known variability in nitrogen-photosynthesis relationships, we develop a mechanistic nitrogen allocation model based on a trade-off of nitrogen allocated between growth and storage, and an optimization of nitrogen allocated among light capture, electron transport, carboxylation, and respiration. The developed model is able to predict the acclimation of photosynthetic capacity to changes in CO2 concentration, temperature, and radiation when evaluated against published data of Vc,max (maximum carboxylation rate) and Jmax (maximum electron transport rate). A sensitivity analysis of the model for herbaceous plants, deciduous and evergreen trees implies that elevated CO2 concentrations lead to lower allocation of nitrogen to carboxylation but higher allocation to storage. Higher growing-season temperatures cause lower allocation of nitrogen to carboxylation, due to higher nitrogen requirements for light capture pigments and for storage. Lower levels of radiation have a much stronger effect on allocation of nitrogen to carboxylation for herbaceous plants than for trees, resulting from higher nitrogen requirements for light capture for herbaceous plants. As far as we know, this is the first model of complete nitrogen allocation that simultaneously considers nitrogen allocation to light capture, electron transport, carboxylation, respiration and storage, and the responses of each to altered environmental conditions. We expect this model could potentially improve our confidence in simulations of carbon-nitrogen interactions and the vegetation feedbacks to climate in Earth system models.
PMCID: PMC3359379  PMID: 22649564
25.  Radial Growth of Two Dominant Montane Conifer Tree Species in Response to Climate Change in North-Central China 
PLoS ONE  2014;9(11):e112537.
North-Central China is a region in which the air temperature has clearly increased for several decades. Picea meyeri and Larix principis-rupprechtii are the most dominant co-occurring tree species within the cold coniferous forest belt ranging vertically from 1800 m to 2800 m a.s.l. in this region. Based on a tree-ring analysis of 292 increment cores sampled from 146 trees at different elevations, this study aimed to examine if the radial growth of the two species in response to climate is similar, whether the responses are consistent along altitudinal gradients and which species might be favored in the future driven by the changing climate. The results indicated the following: (1) The two species grew in different rhythms at low and high elevation respectively; (2) Both species displayed inconsistent relationships between radial growth and climate data along altitudinal gradients. The correlation between radial growth and the monthly mean temperature in the spring or summer changed from negative at low elevation into positive at high elevation, whereas those between the radial growth and the total monthly precipitation displayed a change from positive into negative along the elevation gradient. These indicate the different influences of the horizontal climate and vertical mountainous climate on the radial growth of the two species; (3) The species-dependent different response to climate in radial growth appeared mainly in autumn of the previous year. The radial growth of L. principis-rupprechtii displayed negative responses both to temperature and to precipitation in the previous September, October or November, which was not observed in the radial growth of P. meyeri. (4) The radial growth of both species will tend to be increased at high elevation and limited at low elevation, and L. principis-rupprechtii might be more favored in the future, if the temperature keeps rising.
PMCID: PMC4231039  PMID: 25393738

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