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1.  Xylem and phloem phenology in co-occurring conifers exposed to drought 
Trees (Berlin, Germany : West)  2014;28(4):1161-1171.
The focus of this study was to determine temporal dynamics of xylem and phloem formation in co-occurring deciduous and evergreen coniferous species in a dry inner Alpine environment (750 m a.s.l., Tyrol, Austria). By repeated micro-sampling of the stem, timing of key phenological dates of xylem and phloem formation was compared among mature Pinus sylvestris, Larix decidua and Picea abies during two consecutive years. Xylem formation in P. sylvestris started in mid and late April 2011 and 2012, respectively, and in both years about 2 week later in P. abies and L. decidua. Phloem formation preceded xylem formation on average by 3 week in P. sylvestris, and c. 5 week in P. abies and L. decidua. Based on modeled cell number increase, tracheid production peaked between early through late May 2011 and late May through mid-June 2012. Phloem formation culminated between late April and mid-May in 2011 and in late May 2012. Production of xylem and phloem cells continued for about 4 and 5–6 months, respectively. High variability in xylem increment among years and species is related to exogenous control by climatic factors and species-specific sensitivity to drought, respectively. On the other hand, production of phloem cells was quite homogenous and showed asymptotic decrease with respect to xylem cells indicating endogenous control. Results indicate that onset and culmination of xylem and phloem formation are controlled by early spring temperature, whereby strikingly advanced production of phloem compared to xylem cells suggests lower temperature requirement for initiation of the former.
doi:10.1007/s00468-014-1026-x
PMCID: PMC4110670  PMID: 25071313
Cambium; Drought; Intra-annual radial growth; Phloem formation; Xylogenesis
2.  Xylem and phloem phenology in co-occurring conifers exposed to drought 
Trees (Berlin, Germany : West)  2014;28(4):1161-1171.
Key message
Variability in xylem and phloem phenology among years and species is caused by contrasting temperatures prevailing at the start of the growing season and species-specific sensitivity to drought.
Abstract
The focus of this study was to determine temporal dynamics of xylem and phloem formation in co-occurring deciduous and evergreen coniferous species in a dry inner Alpine environment (750 m a.s.l., Tyrol, Austria). By repeated micro-sampling of the stem, timing of key phenological dates of xylem and phloem formation was compared among mature Pinus sylvestris, Larix decidua and Picea abies during two consecutive years. Xylem formation in P. sylvestris started in mid and late April 2011 and 2012, respectively, and in both years about 2 week later in P. abies and L. decidua. Phloem formation preceded xylem formation on average by 3 week in P. sylvestris, and c. 5 week in P. abies and L. decidua. Based on modeled cell number increase, tracheid production peaked between early through late May 2011 and late May through mid-June 2012. Phloem formation culminated between late April and mid-May in 2011 and in late May 2012. Production of xylem and phloem cells continued for about 4 and 5–6 months, respectively. High variability in xylem increment among years and species is related to exogenous control by climatic factors and species-specific sensitivity to drought, respectively. On the other hand, production of phloem cells was quite homogenous and showed asymptotic decrease with respect to xylem cells indicating endogenous control. Results indicate that onset and culmination of xylem and phloem formation are controlled by early spring temperature, whereby strikingly advanced production of phloem compared to xylem cells suggests lower temperature requirement for initiation of the former.
doi:10.1007/s00468-014-1026-x
PMCID: PMC4110670  PMID: 25071313
Cambium; Drought; Intra-annual radial growth; Phloem formation; Xylogenesis
3.  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
4.  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
5.  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
6.  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

Results 1-6 (6)