Climatically controlled reproduction drives interannual growth variability in a temperate tree species.

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent-wide datasets of tree-ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort ('mast years') is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction

Contrasting controls on tree ring isotope variation for Amazon floodplain and terra firme trees

Isotopes in tropical trees rings can improve our understanding of tree responses to climate. We assessed how climate and growing conditions affect tree-ring oxygen and carbon isotopes (δ18OTR and δ13CTR) in four Amazon trees. We analysed within-ring isotope variation for two terra firme (non-flooded) and two floodplain trees growing at sites with varying seasonality. We find distinct intra-annual patterns of δ18OTR and δ13CTR driven mostly by seasonal variation in weather and source water δ18O. Seasonal variation in isotopes was lowest for the tree growing under the wettest conditions. Tree ring cellulose isotope models based on existing theory reproduced well observed within-ring variation with possible contributions of both stomatal and mesophyll conductance to variation in δ13CTR. Climate analysis reveal that terra firme δ18OTR signals were related to basin-wide precipitation, indicating a source water δ18O influence, while floodplain trees recorded leaf enrichment effects related to local climate. Thus, intrinsically different processes (source water vs leaf enrichment) affect δ18OTR in the two different species analysed. These differences are likely a result of both species-specific traits and of the contrasting growing conditions in the floodplains and terra firme environments. Simultaneous analysis of δ13CTR and δ18OTR supports this interpretation as it shows strongly similar intra-annual patterns for both isotopes in the floodplain trees arising from a common control by leaf stomatal conductance, while terra firme trees showed less covariation between the two isotopes. Our results are interesting from a plant physiological perspective and have implications for climate reconstructions as trees record intrinsically different processes

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and C-13/C-12 in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (delta C-13) of wood. The sensitivity of ring width, wood density and delta C-13 to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July-September). delta C-13 variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by delta C-13 of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season delta C-13 of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and C-13 composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season delta C-13 values of wood between individual trees (-31 to -24 parts per thousand) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930-990 g cm(-3) stop). The relationship between seasonal changes in tree-ring structure and C-13 composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20-30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction. [References: 55

The influence of climate and fructification on the inter-annual variability of stem growth and net primary productivity in an old-growth, mixed beech forest

The periodic production of large seed crops by trees (masting) and its interaction with stern growth has long been the objective of tree physiology research. However, very little is known about the effects of masting on stern growth and total net primary productivity (NPP) at the stand scale. This study was conducted in an old-growth, mixed deciduous forest dominated by Fagus sylvatica (L.) and covers the period from 2003 to 2007, which comprised wet, dry and regular years as well as two masts of Fagus and one mast of the co-dominant tree species Fraxinus excelsior (L.) and Acer pseudoplatanus (L.). We combined analyses of weather conditions and stem growth at the tree level (inter- and intra-annual) with fruit, stem and leaf production, and estimates of total NPP at the stand level. Finally, we compared the annual demand of carbon for biomass production with net canopy assimilation (NCA), derived from eddy covariance flux measurements, chamber measurements and modelling. Annual stem growth of Fagus was most favoured by warm periods in spring and that of Fraxinus by high precipitation in June. For stem growth of Acer and for fruit production, no significant relationships with mean weather conditions were found. Intra-annual stem growth of all species was strongly reduced when the relative plant-available water in soil dropped below a threshold of about 60% between May and July. The inter-annual variations of NCA, total NPP and leaf NPP at the stand level were low (mean values 1313, 662 and 168 g C m(-2) year(-1), respectively), while wood and fruit production varied more and contrarily (wood: 169-241 g C m(-2) year(-1); fruits: 21-142 g C m(-2) year(-1)). In all years, an annual surplus of newly assimilated carbon was calculated (on average 100 g C m(-2) year(-1)). The results suggest that stem growth is generally not limited by insufficient carbon resources; only in mast years a short-term carbon shortage may occur in spring. In contrast to common assumption, stem growth alone is not a sufficient proxy for total biomass production or the control of carbon sequestration by weather extremes

Intra-annual variability of anatomical structure and delta C-13 values within tree rings of spruce and pine inalpine, temperate and boreal Europe

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.ISSN:0029-8549ISSN:1432-193

Intra-annual variability of anatomical structure and delta 13C values within tree rings of spruce and pine in alpine, temperate and boreal Europe

Tree-ring width, wood density, anatomical structure and C-13/C-12 ratios expressed as delta C-13-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, delta C-13 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 delta C-13-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The delta C-13-values were not correlated with precipitation or temperature. A highly significant correlation was also found between delta C-13 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 delta C-13 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 delta C-13 and tree-ring width and climate are multi-factorial in seasonal climates