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A Technical Perspective in Modern Tree-ring Research - How to Overcome Dendroecological and Wood Anatomical Challenges

Dendroecological research uses information stored in tree rings to understand how single trees and even entire forest ecosystems responded to ...

The anatomical features of earlywood vessels often reflect information about past climatic conditions. We examined the relationships between mean monthly temperature and mean vessel lumen area (MVA) in various categories of earlywood vessels. Subsets of earlywood vessels of chestnut (Castanea sativa Mill.) were selected from a previously reported dataset based on several progressive size-related procedures. To include all earlywood vessels, the minimum size considered was 10,000 microm2. Changes in the correlations between MVA and the mean air temperature in March are described and discussed. The results show that not all vessels embody the same information. The MVA of a proportion of the largest earlywood vessels in each annual ring was most closely related to March temperature, whereas MVA of the smallest earlywood vessels was better correlated with June temperature. This difference is probably a result of the vessels being formed at different times: early spring for the largest earlywood vessels and later in spring for the smallest earlywood vessels. Analyses combining large and small vessels yielded lower correlations between MVA and monthly temperature. The number, size and distribution of vessels can vary greatly from ring to ring. In making year-to-year comparisons, the best information is provided by observations on vessels of contemporaneous ontogenesis. Criteria for the selection and analysis of vessels in the assessment of temperature during the season of wood formation are proposed and discussed.

Selecting earlywood vessels to maximize their environmental signal.

The aim of this study was to identify the climatic signal contained in the earlywood vessel size of the ring-porous chestnut (Castanea sativa) and the physiological processes involved in the underlying mechanisms. In order to assign the encoded signal to a specific physiological process, bud phenology and vessel formation were monitored along an elevation transect and chronologies of the size of the first row of earlywood vessels were retrospectively correlated with 40 yr of early spring temperatures. The first vessels appeared in late April to early May, after encoding both a negative temperature signal in February-March (during tree quiescence) and a positive temperature signal in early April (at the time of resumption of shoot growth). We hypothesize that February and March temperatures affect cambial sensitivity to auxin, preconditioning tree responses later in the season. Furthermore, April temperature is related to tree activation whereby new hormone production fosters vessel expansion.

Earlywood vessels of Castanea sativa record temperature before their formation.

Drought impairs tree growth in the inner-Alpine valleys of Central Europe. We investigated species-specific responses to contrasting water supply, with Scots pine (Pinus sylvestris L.), threatened by drought-induced mortality, and pubescent oak (Quercus pubescens Willd.), showing no connection between drought events and mortality. The two co-occurring tree species were compared, growing either along an open water channel or at a site with naturally dry conditions. In addition, the growth response of Scots pine to a draining of a water channel was studied. We analysed the radial increment for the last 100 years and wood anatomical parameters for the last 45 years. Drought reduced the conduit area of pubescent oak, but increased the radial lumen diameter of the conduits in Scots pine. Both species decreased their radial increment under drought. In Scots pine, radial increment was generally more dependent on water availability than that in pubescent oak. Irrigated trees responded less negatively to high temperature as seen in the increase in the conduit area in pubescent oak and the removal of the limitation of cell division by high temperatures. After irrigation stopped, tree-ring width for Scots pine decreased within 1-year delay, whereas lumen diameter and cell-wall thickness responded with a 4-year delay. Scots pine seemed to optimize the carbon-per-conduit-costs under drought by increasing conduits diameter while decreasing cell numbers. This strategy might lead to a complete loss of tree rings under severe drought and thus to an impairment of water transport. In contrast, in pubescent oak tree-ring width is less affected by summer drought because parts of the earlywood are built in early spring. Thus, pubescent oak might have gradual advantages over pine in today's climate of the inner-Alpine valley.

Drought-induced adaptation of the xylem in Scots pine and pubescent oak.

Variability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, sub-annual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that low-cost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.

Studying global change through investigation of the plastic responses of xylem anatomy in tree rings.

The 2007 European larch (Larix decidua Mill.) growing season was monitored along two elevational transects in the Lötschental valley in the Swiss Alps. Phenological observations and weekly microcore sampling of 28 larch trees were conducted between April and October 2007 at seven study sites regularly spaced from 1350 to 2150 m a.s.l. on northwest- and southeast-facing slopes. The developmental stages of nearly 75,000 individual cells assessed on 1200 thin sections were used to investigate the links between the trees' thermal regimes and growth phases including the beginning and ending of cell enlargement, wall thickening and maturation of the stem wood. Needles appeared approximately 3-4 weeks earlier than stem growth. The duration of ring formation lasted from mid-May to the end of October, with the length of the growing season decreasing along elevation from 137 to 101 days. The onset of the different growing seasons changed by 3-4 days per 100 m elevation; the ending of the growing season, however, appeared minimally related to altitude. If associated with the monitored altitudinal lapse rate of -0.5 degrees C per 100 m, these results translate into a lengthening of the growing season by approximately 7 days per degree Celsius. This study provides new data on the timing and duration of basic growth processes and contributes to quantification of the impacts of global warming on tree growth and productivity.

Timing and duration of European larch growing season along altitudinal gradients in the Swiss Alps.

Cambial injury has been reported to alter wood structure in broad-leaved trees. However, the duration and extension of associated anatomical changes have rarely been analysed thoroughly. A total of 18 young European ash (Fraxinus excelsior L.) trees injured on the stem by a spring flood were sampled with the aim of comparing earlywood vessels and rays formed prior to and after the scarring event. Anatomical and hydraulic parameters were measured in five successive rings over one-quarter of the stem circumference. The results demonstrate that mechanical damage induces a decrease in vessel lumen size (up to 77%) and an increase in vessel number (up to 475%) and ray number (up to 115%). The presence of more earlywood vessels and rays was observed over at least three years after stem scarring. By contrast, abnormally narrow earlywood vessels mainly developed in the first ring formed after the event, increasing the thickness-to-span ratio of vessels by 94% and reducing both xylem relative conductivity and the index for xylem vulnerability to cavitation by 54% and 32%, respectively. These vessels accumulated in radial groups in a 30° sector immediately adjacent to the wound, raising the vessel grouping index by 28%. The wound-induced anatomical changes in wood structure express the functional need of trees to improve xylem hydraulic safety and mechanical strength at the expense of water transport. Xylem hydraulic efficiency was restored in one year, while xylem mechanical reinforcement and resistance to cavitation and decay lasted over several years.

Duration and extension of anatomical changes in wood structure after cambial injury.

Vessels of broad-leaved trees have been analyzed to study how trees deal with various environmental factors. Cambial injury, in particular, has been reported to induce the formation of narrower conduits. Yet, little or no effort has been devoted to the elaboration of vessel sampling strategies for retrospective injury detection based on vessel lumen size reduction. To fill this methodological gap, four wounded individuals each of grey alder (Alnus incana (L.) Moench) and downy birch (Betula pubescens Ehrh.) were harvested in an avalanche path. Earlywood vessel lumina were measured and compared for each tree between the injury ring built during the growing season following wounding and the control ring laid down the previous year. Measurements were performed along a 10 mm wide radial strip, located directly next to the injury. Specifically, this study aimed at (i) investigating the intra-annual duration and local extension of vessel narrowing close to the wound margin and (ii) identifying an adequate sample of earlywood vessels (number and intra-ring location of cells) attesting to cambial injury. Based on the results of this study, we recommend analyzing at least 30 vessels in each ring. Within the 10 mm wide segment of the injury ring, wound-induced reduction in vessel lumen size did not fade with increasing radial and tangential distances, but we nevertheless advise favoring early earlywood vessels located closest to the injury. These findings, derived from two species widespread across subarctic, mountainous, and temperate regions, will assist retrospective injury detection in Alnus, Betula, and other diffuse-porous species as well as future related research on hydraulic implications after wounding.

Defining an adequate sample of earlywood vessels for retrospective injury detection in diffuse-porous species.

Xylem plasticity in response to climate.

Xylem structure determines the hydraulic and mechanical properties of a stem, and its plasticity is fundamental for maintaining tree performance under changing conditions. Unveiling the mechanism and the range of xylem adjustment is thus necessary to anticipate climate change impacts on vegetation.

Temperature-induced responses of xylem structure of Larix sibirica (Pinaceae) from the Russian Altay.

Environment and genetics combine to influence tree growth and should therefore be jointly considered when evaluating forest responses in a warming climate. Here, we combine dendroclimatology and population genetic approaches with the aim of attributing climatic influences on growth of European larch (Larix decidua) and Norway spruce (Picea abies). Increment cores and genomic DNA samples were collected from populations along a ~900-m elevational transect where the air temperature gradient encompasses a ~4 °C temperature difference. We found that low genetic differentiation among populations indicates gene flow is high, suggesting that migration rate is high enough to counteract the selective pressures of local environmental variation. We observed lower growth rates towards higher elevations and a transition from negative to positive correlations with growing season temperature upward along the elevational transect. With increasing elevation there was also a clear increase in the explained variance of growth due to summer temperatures. Comparisons between climate sensitivity patterns observed along this elevational transect with those from Larix and Picea sites distributed across the Alps reveal good agreement, and suggest that tree-ring width (TRW) variations are more climate-driven than genetics-driven at regional and larger scales. We conclude that elevational transects are an extremely valuable platform for understanding climatic-driven changes over time and can be especially powerful when working within an assessed genetic framework.

Tree growth response along an elevational gradient: climate or genetics?

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.

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

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.

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

For accurate interpretation of oxygen isotopes in tree rings (δ(18) O), it is necessary to disentangle the mechanisms underlying the variations in the tree's internal water cycle and to understand the transfer of source versus leaf water δ(18) O to phloem sugars and stem wood. We studied the seasonal transfer of oxygen isotopes from precipitation and soil water through the xylem, needles and phloem to the tree rings of Larix decidua at two alpine sites in the Lötschental (Switzerland). Weekly resolved δ(18) O records of precipitation, soil water, xylem and needle water, phloem organic matter and tree rings were developed. Week-to-week variations in needle-water (18) O enrichment were strongly controlled by weather conditions during the growing season. These short-term variations were, however, not significantly fingerprinted in tree-ring δ(18) O. Instead, seasonal trends in tree-ring δ(18) O predominantly mirrored trends in the source water, including recent precipitation and soil water pools. Modelling results support these findings: seasonal tree-ring δ(18) O variations are captured best when the week-to-week variations of the leaf water signal are suppressed. Our results suggest that climate signals in tree-ring δ(18) O variations should be strongest at temperate sites with humid conditions and precipitation maxima during the growing season.

Seasonal transfer of oxygen isotopes from precipitation and soil to the tree ring: source water versus needle water enrichment.

Conifer tree rings are generally composed of large, thin-walled cells of light earlywood followed by narrow, thick-walled cells of dense latewood. Yet, how wood formation processes and the associated kinetics create this typical pattern remains poorly understood. We monitored tree-ring formation weekly over 3 yr in 45 trees of three conifer species in France. Data were used to model cell development kinetics, and to attribute the relative importance of the duration and rate of cell enlargement and cell wall deposition on tree-ring structure. Cell enlargement duration contributed to 75% of changes in cell diameter along the tree rings. Remarkably, the amount of wall material per cell was quite constant along the rings. Consequently, and in contrast with widespread belief, changes in cell wall thickness were not principally attributed to the duration and rate of wall deposition (33%), but rather to the changes in cell size (67%). Cell enlargement duration, as the main driver of cell size and wall thickness, contributed to 56% of wood density variation along the rings. This mechanistic framework now forms the basis for unraveling how environmental stresses trigger deviations (e.g. false rings) from the normal tree-ring structure.

Kinetics of tracheid development explain conifer tree-ring structure.

Responses of alpine tree line ecosystems to increasing atmospheric CO2 concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO2 enrichment (+200 ppm; 2001-2009) and 6 years of soil warming (+4 °C; 2007-2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above-ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO2 enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m(-2) ) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO2 led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above-ground mass was not altered by soil warming or elevated CO2 . However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (-40% for all roots <2 mm in diameter at 0-20 cm soil depth) but no change with CO2 enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.

Article	Year
Selecting earlywood vessels to maximize their environmental signal. Tree physiology\| PubMed ID: 16815831	2006
Earlywood vessels of Castanea sativa record temperature before their formation. The New phytologist\| PubMed ID: 17244050	2007
Drought-induced adaptation of the xylem in Scots pine and pubescent oak. Tree physiology\| PubMed ID: 19483185	2009
Studying global change through investigation of the plastic responses of xylem anatomy in tree rings. The New phytologist\| PubMed ID: 19780986	2010
Timing and duration of European larch growing season along altitudinal gradients in the Swiss Alps. Tree physiology\| PubMed ID: 20008326	2010
Duration and extension of anatomical changes in wood structure after cambial injury. Journal of experimental botany\| PubMed ID: 22378953	2012
Defining an adequate sample of earlywood vessels for retrospective injury detection in diffuse-porous species. PloS one\| PubMed ID: 22761707	2012
Xylem plasticity in response to climate. The New phytologist\| PubMed ID: 22861186	2012
Temperature-induced responses of xylem structure of Larix sibirica (Pinaceae) from the Russian Altay. American journal of botany\| PubMed ID: 23660567	2013
Tree growth response along an elevational gradient: climate or genetics? Oecologia\| PubMed ID: 23771802	2013
Intra-annual dynamics of non-structural carbohydrates in the cambium of mature conifer trees reflects radial growth demands. Tree physiology\| PubMed ID: 24128848	2013
A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere. Annals of botany\| PubMed ID: 24201138	2013
Seasonal transfer of oxygen isotopes from precipitation and soil to the tree ring: source water versus needle water enrichment. The New phytologist\| PubMed ID: 24602089	2014
Kinetics of tracheid development explain conifer tree-ring structure. The New phytologist\| PubMed ID: 24890661	2014
Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation. Global change biology\| PubMed ID: 25471674	2014

Patrick Fonti

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