Intra-annual dynamics of non-structural carbohydrates in the cambium of mature conifer trees reflects radial growth demands

The presence of soluble carbohydrates in the cambial zone, either from sugars recently produced during photosynthesis or from starch remobilized from storage organs, is necessary for radial tree growth. However, considerable uncertainties on carbohydrate dynamics and the consequences on tree productivity exist. This study aims to better understand the variation in different carbon pools at intra-annual resolution by quantifying how cambial zone sugar and starch concentrations fluctuate over the season and in relation to cambial phenology. A comparison between two physiologically different species growing at the same site, i.e., the evergreen Picea abies Karst. and the deciduous Larix decidua Mill., and between L. decidua from two contrasting elevations, is presented to identify mechanisms of growth limitation. Results indicate that the annual cycle of sugar concentration within the cambial zone is coupled to the process of wood formation. The highest sugar concentration is observed when the number of cells in secondary wall formation and lignification stages is at a maximum, subsequent to most radial growth. Starch disappears in winter, while other freeze-resistant non-structural carbohydrates (NSCs) increase. Slight differences in NSC concentration between species are consistent with the differing climate sensitivity of the evergreen and deciduous species investigated. The general absence of differences between elevations suggests that the cambial activity of trees growing at the treeline was not limited by the availability of carbohydrates at the cambial zone but instead by environmental controls on the growing season duration.

The potential of stomata analysis in conifers to estimate presence of conifer trees: examples from the Alps

To estimate whether or not a plant taxon found in the fossil record was locally present may be difficult if only pollen is analyzed. Plant macrofossils, in contrast, provide a clear indication of a taxon's local presence, although in some lake sediments or peats, macrofossils may be rare or degraded. For conifers, the stomata found on pollen slides are derived from needles and thus provide a valuable proxy for local presence and they can be identified to genus level. From previously published studies, a transect across the Alps based on 13 sites is presented. For basal samples in sandy silt above the till with high pollen values of Pinus, for example, we may distinguish pine pollen from distant sources (samples with no stomata), from reworked pollen (samples with stomata present). The first apparent local presence of most conifer genera based on stomata often but not always occurs together with the phase of rapid pollen increase (rational limit). An exception is Larix, with its annual deposition of needles and heavy poorly dispersed pollen, for it often shows the first stomata earlier, at the empirical pollen limit. The decline and potential local extinction of a conifer can sometimes be shown in the stomata record. The decline may have been caused by climatic change, competition, or human impact. In situations where conifers form the timberline, the stomata record may indicate timberline fluctuations. In the discussion of immigration or migration of taxa we advocate the use of the cautious term ``apparent local presence'' to include some uncertainties. Absence of a taxon is impossible to prove.

Causes and consequences of past and projected Scandinavian summer temperatures, 500-2100 AD

Tree rings dominate millennium-long temperature reconstructions and many records originate from Scandinavia, an area for which the relative roles of external forcing and internal variation on climatic changes are, however, not yet fully understood. Here we compile 1,179 series of maximum latewood density measurements from 25 conifer sites in northern Scandinavia, establish a suite of 36 subset chronologies, and analyse their climate signal. A new reconstruction for the 1483–2006 period correlates at 0.80 with June–August temperatures back to 1860. Summer cooling during the early 17th century and peak warming in the 1930s translate into a decadal amplitude of 2.9°C, which agrees with existing Scandinavian tree-ring proxies. Climate model simulations reveal similar amounts of mid to low frequency variability, suggesting that internal ocean-atmosphere feedbacks likely influenced Scandinavian temperatures more than external forcing. Projected 21st century warming under the SRES A2 scenario would, however, exceed the reconstructed temperature envelope of the past 1,500 years.

Rolling stones and tree rings: A state of research on dendrogeomorphic reconstructions of rockfall

This progress report focuses on the contribution of tree-ring series to rockfall research and on recent development and challenges in the field. Dendrogeomorphic techniques have been used extensively since the early 2000s and several approaches have been developed to extract rockfall signals from tree-ring records of conifer trees. The reconstruction of rockfall chronologies has been hampered in the past by sample sizes that decrease as one goes back in time, as well as by a paucity of studies that include broadleaved tree species, which are in fact quite common in rockfall-prone environments. In this report, we propose a new approach considering impact probability and quantification of uncertainty in the reconstruction of rockfall time series as well as a quantitative estimate of presumably missed events. In addition, we outline new approaches and future perspectives for the inclusion of woody vegetation in hazard assessment procedures, and end with future thematic perspectives.

A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere

Background and Aims Ongoing global warming has been implicated in shifting phenological patterns such as the timing and duration of the growing season across a wide variety of ecosystems. Linear models are routinely used to extrapolate these observed shifts in phenology into the future and to estimate changes in associated ecosystem properties such as net primary productivity. Yet, in nature, linear relationships may be special cases. Biological processes frequently follow more complex, non-linear patterns according to limiting factors that generate shifts and discontinuities, or contain thresholds beyond which responses change abruptly. This study investigates to what extent cambium phenology is associated with xylem growth and differentiation across conifer species of the northern hemisphere. Methods Xylem cell production is compared with the periods of cambial activity and cell differentiation assessed on a weekly time scale on histological sections of cambium and wood tissue collected from the stems of nine species in Canada and Europe over 1–9 years per site from 1998 to 2011. Key Results The dynamics of xylogenesis were surprisingly homogeneous among conifer species, although dispersions from the average were obviously observed. Within the range analysed, the relationships between the phenological timings were linear, with several slopes showing values close to or not statistically different from 1. The relationships between the phenological timings and cell production were distinctly non-linear, and involved an exponential pattern. Conclusions The trees adjust their phenological timings according to linear patterns. Thus, shifts of one phenological phase are associated with synchronous and comparable shifts of the successive phases. However, small increases in the duration of xylogenesis could correspond to a substantial increase in cell production. The findings suggest that the length of the growing season and the resulting amount of growth could respond differently to changes in environmental conditions.

New evidence for the symbiosis between Tuber aestivum and Picea abies

The Burgundy truffle (Tuber aestivum Vittad.), an ectomycorrhizal fungus living in association with host plants, is one of the most exclusive delicacies. The symbiosis with deciduous oak, beech, and hazel dominates our concept of truffle ecophysiology, whereas potential conifer hosts have rarely been reported. Here, we present morphological and molecular evidence of a wildlife T. aestivum symbiosis with Norway spruce (Picea abies Karst.) and an independent greenhouse inoculation experiment, to confirm our field observation in southwest Germany. A total of 27 out of 50 P. abies seedlings developed T. aestivum ectomycorrhizae with a mean mycorrhization rate of 19.6 %. These findings not only suggest P. abies to be a productive host species under suitable biogeographic conditions but also emphasize the broad ecological amplitude and great symbiotic range of T. aestivum. While challenging common knowledge, this study demonstrates a significant expansion of the species' cultivation potential to the central European regions, where P. abies forests occur on calcareous soils.

Tracing the origin of Arctic driftwood

Arctic environments, where surface temperatures increase and sea ice cover and permafrost depth decrease, are very sensitive to even slight climatic variations. Placing recent environmental change of the high-northern latitudes in a long-term context is, however, complicated by too short meteorological observations and too few proxy records. Driftwood may represent a unique cross-disciplinary archive at the interface of marine and terrestrial processes. Here, we introduce 1445 driftwood remains from coastal East Greenland and Svalbard. Macroscopy and microscopy were applied for wood anatomical classification; a multi-species subset was used for detecting fungi; and information on boreal vegetation patterns, circumpolar river systems, and ocean current dynamics was reviewed and evaluated. Four conifer (Pinus, Larix, Picea, and Abies) and three deciduous (Populus, Salix, and Betula) genera were differentiated. Species-specific identification also separated Pinus sylvestris and Pinus sibirica, which account for ~40% of all driftwood and predominantly originate from western and central Siberia. Larch and spruce from Siberia or North America represents ~26% and ~18% of all materials, respectively. Fungal colonization caused different levels of driftwood staining and/or decay. Our results demonstrate the importance of combining wood anatomical knowledge with insight on boreal forest composition for successfully tracing the origin of Arctic driftwood. To ultimately reconstruct spatiotemporal variations in ocean currents, and to better quantify postglacial uplift rates, we recommend consideration of dendrochronologically dated material from many more circumpolar sites.

Climatic drivers of hourly to yearly tree radius variations along a 6°C natural warming gradient

Climate affects the timing, rate and dynamics of tree growth, over time scales ranging from seconds to centuries. Monitoring how a tree's stem radius varies over these time scales can provide insight into intra-annual stem dynamics and improve our understanding of climate impacts on tree physiology and growth processes. Here, we quantify the response of radial conifer stem size to environmental fluctuations via a novel assessment of tree circadian cycles. We analyze four years of sub-hourly data collected from 56 larch and spruce trees growing along a natural temperature gradient of ∼6 °C in the central Swiss Alps. During the growing season, tree stem diameters were greatest at mid-morning and smallest in the late evening, reflecting the daily cycle of water uptake and loss. Along the gradient, amplitudes calculated from the stem radius cycle were ∼50% smaller at the upper site (∼2200 m a.s.l.) relative to the lower site (∼800 m a.s.l.). We show changes in precipitation, temperature and cloud cover have a substantial effect on typical growing season diurnal cycles; amplitudes were nine times smaller on rainy days (>10 mm), and daily amplitudes are approximately 40% larger when the mean daily temperature is 15–20 °C than when it is 5–10 °C. We find that over the growing season in the sub-alpine forests, spruce show greater daily stem water movement than larch. However, under projected future warming, larch could experience up to 50% greater stem water use, which may severely affect future growth on already dry sites. Our data further indicate that because of the confounding influences of radial growth and short-term water dynamics on stem size, conventional methodology probably overstates the effect of water-linked meteorological variables (i.e. precipitation and relative humidity) on intra-annual tree growth. We suggest future studies use intra-seasonal measurements of cell development and consider whether climatic factors produce reversible changes in stem diameter. These study design elements may help researchers more accurately quantify and attribute changes in forest productivity in response to future warming.

Site and Age Condition the Growth Responses to Climate and Drought of Relict Pinus nigra subsp. salzmannii Populations in Southern Spain

To assess if tree age may modulate the main climatic drivers of radial growth, two relict Pinus nigra subsp. salzmannii populations (Maria, most xeric site; Magina, least xeric site) were sampled in southern Spain near the limits of the species range. Tree-ring width residual chronologies for two age groups (mature trees, age <= 100 years (minimum 40 years); old trees, age > 100 years) were built to evaluate their responses to climate by relating them to monthly precipitation and temperature and a drought index (DRI) using correlation and response functions. We found that drought is the main driver of growth of relict P. nigra populations, but differences between sites and age classes were also observed. First, growth in the most xeric site depends on the drought severity during the previous autumn and the spring of the year of tree-ring formation, whereas in the relatively more mesic site growth is mainly enhanced by warm and wet conditions in spring. Second, growth of mature trees responded more to drought severity than that of old trees. Our findings indicate that drought severity will mainly affect growth of relict P. nigra populations dominated by mature trees in xeric sites. This conclusion may also apply to similar mountain Mediterranean conifer relicts.

Seven years of annual pollen influx at the forest limit in the Swiss Alps studied by pollen traps: relations to vegetation and climate

Annual pollen influx has been monitored in short transects across the altitudinal tree limit in four areas of the Swiss Alps with the use of modified Tauber traps placed at the ground surface. The study areas are Grindelwald (8 traps), Aletsch (8 traps), Simplon (5 traps), and Zermatt (5 traps). The vegetation around the traps is described. The results obtained are: (1) Peak years of pollen influx (one or two in seven years) follow years of high average air temperatures during June–November of the previous year for Larix and Picea, and less clearly for Pinus non-cembra, but not at all for Pinus cembra and Alnus viridis. (2) At the upper forest limit, the regional pollen influx of trees (trees absent within 100 m of the pollen trap) relates well to the average basal area of the same taxon within 10–15 km of the study areas for Pinus cembra, Larix, and Betula, but not for Picea, Pinus non-cembra, and Alnus viridis. (3) The example of Zermatt shows that pollen influx characterises the upper forest limit, if the latter is more or less intact. (4) Presence/absence of Picea, Pinus cembra, Larix, Pinus non-cembra, and Alnus viridis trees within 50–100 m of the traps is apparent in the pollen influx in peak years of pollen influx but not in other years, suggesting that forest-limit trees produce significant amounts of pollen only in some years. (5) Pollen influx averaged over the study period correlates well with the abundance of plants around the pollen traps for conifer trees (but not deciduous trees), Calluna, Gramineae, and Cyperaceae, and less clearly so Compositae Subfam. Cichorioideae and Potentilla-type. (6) Influx of extra-regional pollen derived from south of the Alps is highest in Simplon, which is open to southerly winds, slightly lower in Aletsch lying just north of Simplon, and lowest in Zermatt sheltered from the south by high mountains and Grindelwald lying north of the central Alps.

Larix decidua and other larches in Europe: distribution, habitat, usage and threats

The European larch (Larix decidua Mill.) is a pioneer, very long-lived, fast-growing coniferous tree, which occurs in the central and eastern mountains of Europe, forming open forests or pasture woods at the upper tree limits. Larch is the only deciduous conifer in Europe as an adaptation to continental alpine climates. In fact, it is able to tolerate very cold temperatures during winter and, by losing its needles, avoids foliage desiccation. It is a transitional species, colonising open terrain after natural disturbances. It forms pure stands but more often it is found with other alpine tree species, which tend to replace it if no other disturbances occur. Thanks to its adaptability and the durability of its wood, the European larch represents an important silvicultural tree species in the alpine regions, planted even outside its natural ranges. Its wood is largely used for carpentry, furniture and pulp for paper. In lower altitudes or with high precipitation rates, larch is more susceptible to fungal diseases. Outbreaks of insect defoliators, principally caused by the larch bud moth (Zeiraphera diniana), can limit tree development, with economic losses in plantations, but they rarely lead to the death of the trees.