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1.  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
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.
doi:10.1007/s00468-007-0166-7
PMCID: PMC3083837  PMID: 21532976
Climate warming; moving response function; Pinus cembra; temperature sensitivity; tree ring
3.  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.
Summary
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.
doi:10.1093/treephys/tpp001
PMCID: PMC3013296  PMID: 19203979
cambial reactivation; dormancy; Pinus cembra; radial stem growth; sap flow; stem CO2 efflux; stem respiration; xylem production
4.  Radial stem growth in response to microclimate and soil moisture in a drought-prone mixed coniferous forest at an inner Alpine site 
European journal of forest research  2014;133(3):467-479.
Dendroclimatological studies in a dry inner Alpine environment (750 m a.s.l.) revealed different growth response of co-occurring coniferous species to climate, which is assumed to be caused by a temporal shift in wood formation among species. The main focus of this study therefore was to monitor intra-annual dynamics of radial increment growth of mature deciduous and evergreen coniferous species (Pinus sylvestris, Larix decidua and Picea abies) during two consecutive years with contrasting climatic conditions. Radial stem growth was continuously followed by band dendrometers and modelled using Gompertz functions to determine time of maximum growth. Histological analyses of tree ring formation allowed determination of temporal dynamics of cambial activity and xylem cell development. Daily fluctuations in stem radius and radial stem increments were extracted from dendrometer traces, and correlations with environmental variables were performed. While a shift in temporal dynamics of radial growth onset and cessation was detected among co-occurring species, intra-annual radial growth peaked synchronously in late May 2011 and early June 2012. Moist atmospheric conditions, i.e. high relative air humidity, low vapour pressure deficit and low air temperature during the main growing period, favoured radial stem increment of all species. Soil water content and soil temperature were not significantly related to radial growth. Although a temporal shift in onset and cessation of wood formation was detected among species, synchronous culmination of radial growth indicates homogenous exogenous and/or endogenous control. The close coupling of radial growth to atmospheric conditions points to the importance of stem water status for intra-annual growth of drought-prone conifers.
doi:10.1007/s10342-013-0777-z
PMCID: PMC4035765  PMID: 24883053
Cambial activity; Climate–growth relationship; Conifers; Dendrometer; Drought; Intra-annual radial growth
5.  Climatic influences on intra-annual stem radial increment of Pinus sylvestris (L.) exposed to drought 
Trees (Berlin, Germany : West)  2010;24(5):887-898.
Within a dry inner Alpine valley in the Eastern Central Alps (750 m a.s.l., Tyrol, Austria) the influence of climate variables (precipitation, air humidity, temperature) and soil water content on intra-annual dynamics of tree-ring development was determined in Scots pine (Pinus sylvestris L.) at two sites differing in soil water availability (xeric and dry-mesic site). Radial stem development was continuously followed during 2007 and 2008 by band dendrometers and repeated micro-sampling of the developing tree rings of mature trees. Daily and seasonal fluctuations of the stem radius, which reached almost half of total annual increment, primarily reflected changes in tree water status and masked radial stem growth especially during drought periods in spring. However, temporal dynamics of intra-annual radial growth determined by both methods were found to be quite similar, when onset of radial growth in dendrometer traces was defined by the occurrence of first enlarging xylem cells. Radial increments during the growing period, which lasted from early April through early August showed statistically significant relationships with precipitation (Kendall τ = 0.234, p < 0.01, and τ = 0.184, p < 0.05, at the xeric and dry-mesic site, respectively) and relative air humidity (Pearson r = 0.290, p < 0.05, and r = 0.306, p < 0.05 at the xeric and dry-mesic site, respectively). Soil water content and air temperature had no influence on radial stem increment. Culmination of radial stem growth was detected at both study plots around mid-May, prior to occurrence of more favourable climatic conditions, i.e. an increase in precipitation during summer. We suggest that the early decrease in radial growth rate is due to a high belowground demand for carbohydrates to ensure adequate resource acquisition on the drought prone substrate.
doi:10.1007/s00468-010-0458-1
PMCID: PMC3191526  PMID: 22003269
Dendrometer; Drought; Dry inner Alpine valley; Pinus sylvestris; Radial growth; Xylem cell analysis
6.  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.
doi:10.1007/s10342-010-0419-7
PMCID: PMC3191523  PMID: 22003357
Non structural carbohydrates; seasonal variation; elevational gradient; timberline ecotone; treeline formation; treelife limitation
7.  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.
Summary
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
8.  Impact of drought on the temporal dynamics of wood formation in Pinus sylvestris 
Tree physiology  2010;30(4):490-501.
Summary
We determined the temporal dynamics of cambial activity and xylem cell differentiation of Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m asl, Tyrol, Austria), where radial growth is strongly limited by drought in spring. Repeated micro-sampling of the developing tree ring of mature trees was carried out during 2 contrasting years at two study plots that differ in soil water availability (xeric and dry-mesic site).
In 2007, when air temperature at the beginning of the growing season in April exceeded the long-term mean by 6.4 °C, cambial cell division started in early April at both study plots. A delayed onset of cambial activity of c. 2 wk was found in 2008, when average climate conditions prevailed in spring, indicating that resumption of cambial cell division after winter dormancy is temperature-controlled. Cambial cell division consistently ended about the end of June/early July in both study years. Radial enlargement of tracheids started almost 3 wk earlier in 2007 compared with 2008 at both study plots. At the xeric site, the maximum rate of tracheid production in 2007 and 2008 was reached in early and mid-May, respectively, and c. 2 wk later, at the dry-mesic site. Since in both study years, more favorable growing conditions (i.e., an increase in soil water content) were recorded during summer, we suggest a strong sink competition for carbohydrates to mycorrhizal root and shoot growth. Wood formation stopped c. 4 wk earlier at the xeric compared with the dry-mesic site in both years, indicating a strong influence of drought stress on cell differentiation. This is supported by radial widths of earlywood cells, which were found to be significantly narrower at the xeric than at the dry-mesic site (P < 0.05).
Repeated cellular analyses during the two growing seasons revealed that, although spatial variability in the dynamics and duration of cell differentiation processes in Pinus sylvestris exposed to drought is strongly influenced by water availability, the onset of cambial activity and cell differentiation is controlled by temperature.
doi:10.1093/treephys/tpq003
PMCID: PMC3046340  PMID: 20197285
Cambium; dry inner Alpine valley; intra-annual growth; Scots pine; tracheid production; xylogenesis
9.  Radial Growth of Qilian Juniper on the Northeast Tibetan Plateau and Potential Climate Associations 
PLoS ONE  2013;8(11):e79362.
There is controversy regarding the limiting climatic factor for tree radial growth at the alpine treeline on the northeastern Tibetan Plateau. In this study, we collected 594 increment cores from 331 trees, grouped within four altitude belts spanning the range 3550 to 4020 m.a.s.l. on a single hillside. We have developed four equivalent ring-width chronologies and shown that there are no significant differences in their growth-climate responses during 1956 to 2011 or in their longer-term growth patterns during the period AD 1110–2011. The main climate influence on radial growth is shown to be precipitation variability. Missing ring analysis shows that tree radial growth at the uppermost treeline location is more sensitive to climate variation than that at other elevations, and poor tree radial growth is particularly linked to the occurrence of serious drought events. Hence water limitation, rather than temperature stress, plays the pivotal role in controlling the radial growth of Sabina przewalskii Kom. at the treeline in this region. This finding contradicts any generalisation that tree-ring chronologies from high-elevation treeline environments are mostly indicators of temperature changes.
doi:10.1371/journal.pone.0079362
PMCID: PMC3828375  PMID: 24244488
10.  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.
doi:10.1051/forest/2009023
PMCID: PMC3047780  PMID: 21379395
Alpine timberline; treeline; global warming; CO2; ozone; water balance; carbon gain
11.  Using Automated Point Dendrometers to Analyze Tropical Treeline Stem Growth at Nevado de Colima, Mexico 
Sensors (Basel, Switzerland)  2010;10(6):5827-5844.
The relationship between wood growth and environmental variability at the tropical treeline of North America was investigated using automated, solar-powered sensors (a meteorological station and two dendrometer clusters) installed on Nevado de Colima, Mexico (19° 35′ N, 103° 37′ W, 3,760 m a.s.l.). Pure stands of Pinus hartwegii Lindl. (Mexican mountain pine) were targeted because of their suitability for tree-ring analysis in low-latitude, high-elevation, North American Monsoon environments. Stem size and hydroclimatic variables recorded at half-hour intervals were summarized on a daily timescale. Power outages, insect outbreaks, and sensor failures limited the analysis to non-consecutive months during 2001–2003 at one dendrometer site, and during 2002–2005 at the other. Combined data from the two sites showed that maximum radial growth rates occur in late spring (May), as soil temperature increases, and incoming short-wave radiation reaches its highest values. Early season (April–May) radial increment correlated directly with temperature, especially of the soil, and with solar radiation. Stem expansion at the start of the summer monsoon (June–July) was mostly influenced by moisture, and revealed a drought signal, while late season relationships were more varied.
doi:10.3390/s100605827
PMCID: PMC3247734  PMID: 22219689
point dendrometers; radial growth; tree rings; dendroecology; high elevation ecosystems; Mexican mountain pine; Pinus hartwegii Lindl.; Volcán de Fuego
12.  Effects of environmental conditions on onset of xylem growth in Pinus sylvestris under drought 
Tree physiology  2011;31(5):483-493.
Summary
We determined influence of environmental factors (air and soil temperature, precipitation, photoperiod) on onset of xylem growth in Scots pine (Pinus sylvestris L.) within a dry inner Alpine valley (750 m a.s.l., Tyrol, Austria) by repeatedly sampling micro-cores throughout 2007-2010 at two sites (xeric and dry-mesic) at the start of the growing season. Temperature sums were calculated in degree-days (DD) ≥ 5 °C from 1 January and 20 March, i.e. spring equinox, to account for photoperiodic control of release from winter dormancy. Threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regression. Onset of xylem growth, which was not significantly different between the xeric and dry-mesic site, ranged from mid-April in 2007 to early May in 2008. Among most study years statistically significant differences (P < 0.05) in onset of xylem growth were detected. Mean air temperature sums calculated from 1 January until onset of xylem growth were 230 ± 44 DD (mean ± standard deviation) at the xeric and 205 ± 36 DD at the dry-mesic site. Temperature sums calculated from spring equinox until onset of xylem growth showed quite less variability during the four year study period amounting to 144 ± 10 and 137 ± 12 DD at the xeric and dry-mesic site, respectively. At both sites xylem growth was active when daily minimum, mean and maximum air temperatures were 5.3, 10.1 and 16.2 °C, respectively. Soil temperature thresholds and DD until onset of xylem growth differed significantly between sites indicating minor importance of root-zone temperature for onset of xylem growth. Although spring precipitation is known to limit radial growth in P. sylvestris exposed to dry inner Alpine climate, results of this study revealed that (i) a daily minimum air temperature threshold for onset of xylem growth in the range of 5-6 °C exists and (ii) air temperature sum rather than precipitation or soil temperature triggers start of xylem growth. Based on these findings we suggest that drought stress forces P. sylvestris to draw upon water reserves in the stem for enlargement of first tracheids after cambial resumption in spring.
doi:10.1093/treephys/tpr034
PMCID: PMC3427020  PMID: 21593011
dry inner Alpine valley; heat-sum; phenology; Scots pine; wood formation; xylogenesis
13.  Climate change and the northern Russian treeline zone 
The Russian treeline is a dynamic ecotone typified by steep gradients in summer temperature and regionally variable gradients in albedo and heat flux. The location of the treeline is largely controlled by summer temperatures and growing season length. Temperatures have responded strongly to twentieth-century global warming and will display a magnified response to future warming. Dendroecological studies indicate enhanced conifer recruitment during the twentieth century. However, conifers have not yet recolonized many areas where trees were present during the Medieval Warm period (ca AD 800–1300) or the Holocene Thermal Maximum (HTM; ca 10 000–3000 years ago). Reconstruction of tree distributions during the HTM suggests that the future position of the treeline due to global warming may approximate its former Holocene maximum position. An increased dominance of evergreen tree species in the northern Siberian forests may be an important difference between past and future conditions. Based on the slow rates of treeline expansion observed during the twentieth century, the presence of steep climatic gradients associated with the current Arctic coastline and the prevalence of organic soils, it is possible that rates of treeline expansion will be regionally variable and transient forest communities with species abundances different from today's may develop.
doi:10.1098/rstb.2007.2200
PMCID: PMC2606780  PMID: 18006415
Arctic; Eurasia; Holocene; boreal forest; treeline; climate change
14.  Growth and Phenology of Three Dwarf Shrub Species in a Six-Year Soil Warming Experiment at the Alpine Treeline 
PLoS ONE  2014;9(6):e100577.
Global warming can have substantial impacts on the phenological and growth patterns of alpine and Arctic species, resulting in shifts in plant community composition and ecosystem dynamics. We evaluated the effects of a six-year experimental soil warming treatment (+4°C, 2007–2012) on the phenology and growth of three co-dominant dwarf shrub species growing in the understory of Larix decidua and Pinus uncinata at treeline in the Swiss Alps. We monitored vegetative and reproductive phenology of Vaccinium myrtillus, Vaccinium gaultherioides and Empetrum hermaphroditum throughout the early growing season of 2012 and, following a major harvest at peak season, we measured the biomass of above-ground ramet fractions. For all six years of soil warming we measured annual shoot growth of the three species and analyzed ramet age and xylem ring width of V. myrtillus. Our results show that phenology of the three species was more influenced by snowmelt timing, and also by plot tree species (Larix or Pinus) in the case of V. myrtillus, than by soil warming. However, the warming treatment led to increased V. myrtillus total above-ground ramet biomass (+36% in 2012), especially new shoot biomass (+63% in 2012), as well as increased new shoot increment length and xylem ring width (+22% and +41%, respectively; average for 2007–2012). These results indicate enhanced overall growth of V. myrtillus under soil warming that was sustained over six years and was not caused by an extended growing period in early summer. In contrast, E. hermaphroditum only showed a positive shoot growth response to warming in 2011 (+21%), and V. gaultherioides showed no significant growth response. Our results indicate that V. myrtillus might have a competitive advantage over the less responsive co-occurring dwarf shrub species under future global warming.
doi:10.1371/journal.pone.0100577
PMCID: PMC4067323  PMID: 24956273
15.  Changes in ground beetle assemblages above and below the treeline of the Dolomites after almost 30 years (1980/2009) 
Ecology and Evolution  2014;4(8):1284-1294.
Very little is known about the changes of ground beetle assemblages in the last few decades in the Alps, and different responses to climate change of animal populations living above and below the treeline have not been estimated yet. This study focuses on an altitudinal habitat sequence from subalpine spruce forest to alpine grassland in a low disturbance area of the southeastern Dolomites in Italy, the Paneveggio Regional Park. We compared the ground beetle (Carabidae) populations sampled in 1980 in six stands below and above the treeline (1650–2250 m a.s.l.) with those sampled in the same sites almost 30 years later (2008/9). Quantitative data (species richness and abundance) have been compared by means of several diversity indexes and with a new index, the Index of Rank-abundance Change (IRC). Our work shows that species richness and abundance have changed after almost 30 years as a consequence of local extinctions, uphill increment of abundance and uphill shift of distribution range. The overall species number dropped from 36 to 27, while in the sites above the treeline, species richness and abundance changed more than in the forest sites. Two microtherm characteristic species of the pioneer cushion grass mats, Nebria germari and Trechus dolomitanus, became extinct or showed strong abundance reduction. In Nardetum pastures, several hygrophilic species disappeared, and xerophilic zoophytophagous elements raised their population density. In forest ecosystems, the precipitation reduction caused deep soil texture and watering changes, driving a transformation from Sphagnum-rich (peaty) to humus-rich soil, and as a consequence, soil invertebrate biomass strongly increased and thermophilic carabids enriched the species structure. In three decades, Carabid assemblages changed consistently with the hypothesis that climate change is one of the main factors triggering natural environment modifications. Furthermore, the level of human disturbance could enhance the sensitivity of mountain ecosystems to climate change.
doi:10.1002/ece3.927
PMCID: PMC4020689  PMID: 24834326
Carabids; dolomites; global change; mountain
16.  Climate Control on Tree Growth at the Upper and Lower Treelines: A Case Study in the Qilian Mountains, Tibetan Plateau 
PLoS ONE  2013;8(7):e69065.
It is generally hypothesized that tree growth at the upper treeline is normally controlled by temperature while that at the lower treeline is precipitation limited. However, uniform patterns of inter-annual ring-width variations along altitudinal gradients are also observed in some situations. How changing elevation influences tree growth in the cold and arid Qilian Mountains, on the northeastern Tibetan Plateau, is of considerable interest because of the sensitivity of the region’s local climate to different atmospheric circulation patterns. Here, a network of four Qilian juniper (Sabina przewalskii Kom.) ring-width chronologies was developed from trees distributed on a typical mountain slope at elevations ranging from 3000 to 3520 m above sea level (a.s.l.). The statistical characteristics of the four tree-ring chronologies show no significant correlation with increasing elevation. All the sampled tree growth was controlled by a common climatic signal (local precipitation) across the investigated altitudinal gradient (520 m). During the common reliable period, covering the past 450 years, the four chronologies have exhibited coherent growth patterns in both the high- and low-frequency domains. These results contradict the notion of contrasting climate growth controls at higher and lower elevations, and specifically the assumption that inter-annual tree-growth variability is controlled by temperature at the upper treeline. It should be stressed that these results relate to the relatively arid conditions at the sampling sites in the Qilian Mountains.
doi:10.1371/journal.pone.0069065
PMCID: PMC3708892  PMID: 23874871
17.  Climate Warming and the Recent Treeline Shift in the European Alps: The Role of Geomorphological Factors in High-Altitude Sites 
Ambio  2010;40(3):264-273.
Global warming and the stronger regional temperature trends recently recorded over the European Alps have triggered several biological and physical dynamics in high-altitude environments. We defined the present treeline altitude in three valleys of a region in the western Italian Alps and reconstructed the past treeline position for the last three centuries in a nearly undisturbed site by means of a dendrochronological approach. We found that the treeline altitude in this region is mainly controlled by human impacts and geomorphological factors. The reconstruction of the altitudinal dynamics at the study site reveals that the treeline shifted upwards of 115 m over the period 1901–2000, reaching the altitude of 2505 m in 2000 and 2515 m in 2008. The recent treeline shift and the acceleration of tree colonization rates in the alpine belt can be mainly ascribed to the climatic input. However, we point out the increasing role of geomorphological factors in controlling the future treeline position and colonization patterns in high mountains.
doi:10.1007/s13280-010-0096-2
PMCID: PMC3357808  PMID: 21644455
Climate change; Treeline; Geomorphology; Tree rings; Larix decidua; European Alps
18.  Cambial activity related to tree size in a mature silver-fir plantation 
Annals of Botany  2011;108(3):429-438.
Background and Aims
Our knowledge about the influences of environmental factors on tree growth is principally based on the study of dominant trees. However, tree social status may influence intra-annual dynamics of growth, leading to differential responses to environmental conditions. The aim was to determine whether within-stand differences in stem diameters of trees belonging to different crown classes resulted from variations in the length of the growing period or in the rate of cell production.
Methods
Cambial activity was monitored weekly in 2006 for three crown classes in a 40-year-old silver-fir (Abies alba) plantation near Nancy (France). Timings, duration and rate of tracheid production were assessed from anatomical observations of the developing xylem.
Key Results
Cambial activity started earlier, stopped later and lasted longer in dominant trees than in intermediate and suppressed ones. The onset of cambial activity was estimated to have taken 3 weeks to spread to 90 % of the trees in the stand, while the cessation needed 6 weeks. Cambial activity was more intense in dominant trees than in intermediate and suppressed ones. It was estimated that about 75 % of tree-ring width variability was attributable to the rate of cell production and only 25 % to its duration. Moreover, growth duration was correlated to tree height, while growth rate was better correlated to crown area.
Conclusions
These results show that, in a closed conifer forest, stem diameter variations resulted principally from differences in the rate of xylem cell production rather than in its duration. Tree size interacts with environmental factors to control the timings, duration and rate of cambial activity through functional processes involving source–sink relationships principally, but also hormonal controls.
doi:10.1093/aob/mcr168
PMCID: PMC3158687  PMID: 21816842
Cambial activity; forest-stand structure; silver fir (Abies alba); tree-ring formation; tree-to-tree competition; social status; wood anatomy; xylem cell differentiation
19.  Photosynthetic temperature adaptation of Pinus cembra within the timberline ecotone of the Central Austrian Alps 
Annals of forest science  2010;67(2):201.
Temperature is suggested to determine the upper limit of tree life. Therefore, future climate warming may be of importance for tree distribution within the European Alps, where low temperatures limit carbon metabolism.
We focused on the effects of air and soil temperature on net photosynthesis (Pn) of Pinus cembra an evergreen climax species of the timberline ecotone of the Central Austrian Alps. Light response and temperature response curves were estimated along an altitudinal gradient ranging from the forest limit up to the krummholz limit in both summer and fall.
In general, Pn was significantly lower in fall as compared to summer. Nevertheless, independent from season mean Pn values tended to increase with elevation and were positively correlated with root zone temperatures. The specific leaf area by contrast declined with increasing elevation. Furthermore, the temperature optimum of net photosynthesis declined with increasing elevation and was positively correlated with the mean maximum air temperature of the 10 days prior the date of measurement.
Thus, our findings appear to reflect a long-term adaptation of the photosynthetic apparatus of Pinus cembra to the general temperature conditions with respect to elevation combined with a short term acclimation to the prevailing temperature regime.
doi:10.1051/forest/2009094
PMCID: PMC3047779  PMID: 21379394
net photosynthesis; temperature; cembran pine; timberline ecotone; global warming
20.  Elevational patterns of Polylepis tree height (Rosaceae) in the high Andes of Peru: role of human impact and climatic conditions 
We studied tree height in stands of high-Andean Polylepis forests in two cordilleras near Cuzco (Peru) with respect to variations in human impact and climatic conditions, and compared air and soil temperatures between qualitatively defined dry and humid slopes. We studied 46 forest plots of 100 m2 of five Polylepis species at 3560–4680 m. We measured diameter at breast height (dbh) and tree height in the stands (1229 trees in total), as well as air and soil temperatures in a subset of plots. The data was analyzed combining plots of given species from different sites at the same elevation (±100 m). There was no elevational decrease of mean maximum tree height across the entire data set. On humid slopes, tree height decreased continuously with elevation, whereas on dry slopes it peaked at middle elevations. With mean maximum tree heights of 9 m at 4530 m on the humid slopes and of 13 m at 4650 m on the dry slopes, we here document the tallest high-elevation forests found so far worldwide. These highest stands grow under cold mean growing season air temperatures (3.6 and 3.8°C on humid vs. dry slopes) and mean growing season soil temperatures (5.1 vs. 4.6°C). Mean annual air and soil temperature both decreased with elevation. Dry slopes had higher mean and maximum growing season air temperatures than humid slopes. Mean annual soil temperatures did not significantly differ and mean annual air temperatures only slightly differed between slopes. However, maximum air temperatures differed on average by 6.6 K between dry and humid slopes. This suggests that the differences in tree height between the two slopes are most likely due to differences in solar radiation as reflected by maximum air temperatures. Our study furthermore provides evidence that alpine Polylepis treelines grow under lower temperature conditions than global high-elevation treelines on average, suggesting that Polylepis species may have evolved special physiological adaptations to low temperatures.
doi:10.3389/fpls.2014.00194
PMCID: PMC4023064  PMID: 24847343
air temperatures; forest structure; soil temperatures; solar radiation; alpine treeline; tropical forest
21.  A Patchy Growth via Successive and Simultaneous Cambia: Key to Success of the Most Widespread Mangrove Species Avicennia marina? 
Annals of Botany  2007;101(1):49-58.
Background and Aims
Secondary growth via successive cambia has been intriguing researchers for decades. Insight into the mechanism of growth layer formation is, however, limited to the cellular level. The present study aims to clarify secondary growth via successive cambia in the mangrove species Avicennia marina on a macroscopic level, addressing the formation of the growth layer network as a whole. In addition, previously suggested effects of salinity on growth layer formation were reconsidered.
Methods
A 1-year cambial marking experiment was performed on 80 trees from eight sites in two mangrove forests in Kenya. Environmental (soil water salinity and nutrients, soil texture, inundation frequency) and tree characteristics (diameter, height, leaf area index) were recorded for each site. Both groups of variables were analysed in relation to annual number of growth layers, annual radial increment and average growth layer width of stem discs.
Key Results
Between trees of the same site, the number of growth layers formed during the 1-year study period varied from only part of a growth layer up to four growth layers, and was highly correlated to the corresponding radial increment (0–5 mm year–1), even along the different sides of asymmetric stem discs. The radial increment was unrelated to salinity, but the growth layer width decreased with increasing salinity and decreasing tree height.
Conclusions
A patchy growth mechanism was proposed, with an optimal growth at distinct moments in time at different positions around the stem circumference. This strategy creates the opportunity to form several growth layers simultaneously, as observed in 14 % of the studied trees, which may optimize tree growth under favourable conditions. Strong evidence was provided for a mainly endogenous trigger controlling cambium differentiation, with an additional influence of current environmental conditions in a trade-off between hydraulic efficiency and mechanical stability.
doi:10.1093/aob/mcm280
PMCID: PMC2701843  PMID: 18006508
Avicenia marina; cambial marking; mangrove; phloem; salinity; secondary growth; successive cambia; xylem
22.  Effect of Local Heating and Cooling on Cambial Activity and Cell Differentiation in the Stem of Norway Spruce (Picea abies) 
Annals of Botany  2006;97(6):943-951.
• Background and Aims The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated.
• Methods A heating experiment (23–25 °C) was carried out in spring, before normal reactivation of the cambium, and cooling (9–11 °C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees.
• Key Results Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples.
• Conclusions Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.
doi:10.1093/aob/mcl050
PMCID: PMC2803384  PMID: 16613904
Norway spruce; Picea abies; cambium; xylem; phloem; cell differentiation; heating; cooling; light microscopy; transmission electron microscopy; UV-microspectrophotometry
23.  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.
Methods
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.
Conclusions
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.
doi:10.1093/aob/mcq237
PMCID: PMC3025735  PMID: 21186240
Abies georgei; biomechanics; tensile resistance; cellulose content; non-structural carbon; root strength; Tibet–Qinghai; slope stability
24.  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.
doi:10.1007/s00468-012-0768-6
PMCID: PMC3750198  PMID: 23976821
Basal area increment; Dendroclimatology; Inner Alpine valley; Radial growth; Moving response function; Tree-ring analysis
25.  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.
doi:10.1007/s00442-009-1421-y
PMCID: PMC2744769  PMID: 19653008
Dendrochonology; Carbohydrate storage; Picea abies; Pinus cembra; Pinus sylvestris; Tracheid lumen area; Wood density

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