Purpose of Review
Society is concerned about the long-term condition of the forests. Although a clear definition of forest health is still missing, to evaluate forest health, monitoring efforts in ...the past 40 years have concentrated on the assessment of tree vitality, trying to estimate tree photosynthesis rates and productivity. Used in monitoring forest decline in Central Europe since the 1980s, crown foliage transparency has been commonly believed to be the best indicator of tree condition in relation to air pollution, although annual variations appear more closely related to water stress. Although crown transparency is not a good indicator of tree photosynthesis rates, defoliation is still one of the most used indicators of tree vitality. Tree rings have been often used as indicators of past productivity. However, long-term tree growth trends are difficult to interpret because of sampling bias, and ring width patterns do not provide any information about tree physiological processes.
Recent Findings
In the past two decades, tree-ring stable isotopes have been used not only to reconstruct the impact of past climatic events, such as drought, but also in the study of forest decline induced by air pollution episodes, and other natural disturbances and environmental stress, such as pest outbreaks and wildfires. They have proven to be useful tools for understanding physiological processes and tree response to such stress factors.
Summary
Tree-ring stable isotopes integrate crown transpiration rates and photosynthesis rates and may enhance our understanding of tree vitality. They are promising indicators of tree vitality. We call for the use of tree-ring stable isotopes in future monitoring programmes.
Treeline shift and tree growth often respond to climatic changes and it is critical to identify and quantify their dynamics. Some regions are particularly sensitive to climate change and the Altai ...Mountains, located in Central and East Asia, are showing unequivocal signs. The mean annual temperature in the area has increased by 1.3-1.7 °C in the last century. As this mountain range has ancient and protected forests on alpine slopes, we focus on determining the treeline structure and dynamics. We integrated in situ fine-scale allometric data with analyses from dendrochronological samples, high-resolution 3D drone photos and new satellite images to study the dynamics and underlying causal mechanisms of any treeline movement and growth changes in a remote preserved forest at the Aktru Research Station in the Altai Mountain. We show that temperature increase has a negative effect on mountain tree growth. In contrast, only younger trees grow at higher altitudes and we document a relatively fast upward shift of the treeline. During the last 52 years, treeline moved about 150 m upward and the rate of movement accelerated until recently. Before the 1950s, it never shifted over 2150-2200 m a.s.l. We suggest that a continuous upward expansion of the treeline would be at the expense of meadow and shrub species and radically change this high-mountain ecosystem with its endemic flora. This documented treeline shift represents clear evidence of the increased velocity of climate change during the last century.
Summary
Atmospheric carbon dioxide concentration (CO2) is increasing, which increases leaf‐scale photosynthesis and intrinsic water‐use efficiency. These direct responses have the potential to ...increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of CO2 increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a CO2‐driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing CO2 (iCO2) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre‐industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2, albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.
See also the Commentary on this article by Way et al., 229: 2383–2385.
Two simplifying hypotheses have been proposed for whole‐plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first‐principles carbon balance model with a ...prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carry‐over of fixed carbon between years, while the second implies far too great an increase in respiration during stand development—leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.
Metabolism, reserves and size all drive forest C‐balance.
The occurrence of old-growth forests is quite limited in Mediterranean islands, which have been subject to particularly pronounced human impacts. Little is known about the carbon stocks of such ...peculiar ecosystems compared with different stages of secondary succession. We investigated the carbon variation in aboveground woody biomass, in litter and soil, and the nitrogen variation in litter and soil, in a 100 years long secondary succession in Mediterranean ecosystems. A vineyard, three stages of plant succession (high maquis, maquis-forest, and forest-maquis), and an old growth forest were compared. Soil samples at two soil depths (0-15 and 15-30 cm), and two litter types, relatively undecomposed and partly decomposed, were collected. Carbon stock in aboveground woody biomass increased from 6 Mg ha-1 in the vineyard to 105 Mg ha-1 in old growth forest. Along the secondary succession, soil carbon considerably increased from about 33 Mg ha-1 in the vineyard to about 69 Mg ha-1 in old growth forest. Soil nitrogen has more than doubled, ranging from 4.1 Mg ha-1 in the vineyard to 8.8 Mg ha-1 in old growth forest. Both soil parameters were found to be affected by successional stage and soil depth but not by their interaction. While the C/N ratio in the soil remained relatively constant during the succession, the C/N ratio of the litter strongly decreased, probably following the progressive increase in the holm oak contribution. While carbon content in litter decreased along the succession, nitrogen content slightly increased. Overall, carbon stock in aboveground woody biomass, litter and soil increased from about 48 Mg ha-1 in the vineyard to about 198 Mg ha-1 in old growth forest. The results of this study indicate that, even in Mediterranean environments, considerable amounts of carbon may be stored through secondary succession processes up to old growth forest.
•Three thinning treatments of different intensity were applied to Quercus robur mixed plantation.•Tree growth and WUEi were affected by crown asymmetry and competition.•High presence of alder ...influenced tree growth due to resources competitions.•The very high selective thinning resulted to be the most appropriate management practice.•Thinning increased productivity and limited water loss.
Thinning intensity is an important factor affecting forest structure and density and influencing tree growth.
The main objective of the study is to evaluate how the different intensity of thinning can influence, in the short and long term, the productivity and intrinsic water use efficiency (WUEi) of Quercus robur L., a widespread European species. Three thinning types of different intensity were applied in 14-year mixed plantation of Quercus robur L. growing with Alnus glutinosa (L) Gaertn., a N-fixing species, with an initial density of 625 trees/ha. A multidisciplinary approach was applied, coupling the tree-ring analyses with the measurements of stable isotopes of carbon and oxygen.
Findings highlighted how the different conditions of light and competition, determined by the type and intensity of thinning, triggered different responses of Q. robur species in terms of growth and WUEi.
Data suggested that the moderate selective thinning (31% reduction of total biomass) was not able to reduce the high competition for resources among individuals: this condition negatively influenced the trees growth. The presence of the N-fixing species, left after the thinning, was not able to counteract the negative effect of competition.
A negative effect on productivity was recorded with the geometric thinning (62% reduction of total biomass). Indeed, this treatment resulted in an increase of asymmetry of the canopy (0.90 vs 0.83, respectively in pre- and post-thinning period), due to the partial crown competition from one side and with a consequent limitation of the photosynthetic activity of the trees. Finally, the very high selective thinning (75% reduction of total biomass) resulted to be the most appropriate management practice, as it guarantees positive effects both in terms of growth and intrinsic water use efficiency. The post thinning conditions determined high photosynthesis rates, an increase of productivity and a limited water loss.
For accurate interpretation of oxygen isotopes in tree rings (δ¹⁸O), 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 δ¹⁸O 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 δ¹⁸O records of precipitation, soil water, xylem and needle water, phloem organic matter and tree rings were developed. Week‐to‐week variations in needle‐water ¹⁸O enrichment were strongly controlled by weather conditions during the growing season. These short‐term variations were, however, not significantly fingerprinted in tree‐ring δ¹⁸O. Instead, seasonal trends in tree‐ring δ¹⁸O predominantly mirrored trends in the source water, including recent precipitation and soil water pools. Modelling results support these findings: seasonal tree‐ring δ¹⁸O 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 δ¹⁸O variations should be strongest at temperate sites with humid conditions and precipitation maxima during the growing season.
Elevated CO2 increases intrinsic water use efficiency (WUE
i
) of forests, but the magnitude of this effect and its interaction with climate is still poorly understood.
We combined tree ring analysis ...with isotope measurements at three Free Air CO2 Enrichment (FACE, POP-EUROFACE, in Italy; Duke FACE in North Carolina and ORNL in Tennes-see, USA) sites, to cover the entire life of the trees. We used δ13C to assess carbon isotope discrimination and changes in water-use efficiency, while direct CO2 effects on stomatal conductance were explored using δ18O as a proxy.
Across all the sites, elevated CO2 increased 13C-derived water-use efficiency on average by 73% for Liquidambar styraciflua, 77% for Pinus taeda and 75% for Populus sp., but through different ecophysiological mechanisms.
Our findings provide a robust means of predicting water-use efficiency responses from a variety of tree species exposed to variable environmental conditions over time, and species- specific relationships that can help modelling elevated CO2 and climate impacts on forest productivity, carbon and water balances.
Tree mortality and forest dieback episodes are increasing due to drought and heat stress. Nevertheless, a comprehensive understanding of mechanisms enabling trees to withstand and survive droughts ...remains lacking. Our study investigated basal area increment (BAI), and δ
C-derived intrinsic water-use-efficiency (
WUE), to elucidate beech resilience across four healthy stands in Italy with varying climates and soil water availability. Additionally, fist-order autocorrelation (AR1) analysis was performed to detect early warning signals for potential tree dieback risks during extreme drought events. Results reveal a negative link between BAI and vapour pressure deficit (VPD), especially in southern latitudes. After the 2003 drought, BAI decreased at the northern site, with an increase in δ
C and
WUE, indicating conservative water-use. Conversely, the southern sites showed increased BAI and
WUE, likely influenced by rising CO
and improved water availability. In contrast, the central site sustained higher transpiration rates due to higher soil water holding capacity (SWHC). Despite varied responses, most sites exhibited reduced resilience to future extreme events, indicated by increased AR1. Temperature significantly affected beech
WUE and BAI in northern Italy, while VPD strongly influenced the southern latitudes. The observed increase in BAI and
WUE in southern regions might be attributed to an acclimation response.
The effect of fire severity on
growth and ecophysiological responses was evaluated in four burned sites of Vesuvio National Park, Southern Italy. After the wildfire of 2017, when over 1300 hectares ...of vegetation, mainly
woods, were destroyed, four sites were selected according to the different degree of fire severity and a multidisciplinary approach based on tree rings, stable isotopes and percentage of crown scorched or consumed was applied. All the sampled trees in the burned sites showed a decrease in tree growth in 2017, in particular in the latewood at high-severity site. The dendrochronology analyses showed that several individuals experienced and endured higher fire severity in the past compared to 2017 fire. Further δ
C and δ
O underlined the ecophysiological responses and recovery mechanisms of
, suggesting a drastic reduction of photosynthetic and stomata activity in the year of the fire. Our findings demonstrated that
growth reduction is strictly linked to the percentage of crown scorch and that even trees with high level of crown scorched could survive. In all the burned sites the high temperatures and the time of exposure to the flames were not sufficient to determine the death of the cambium and all the trees were able to complete the 2017 seasonal wood formation. This data can contribute to define guidelines to managers making post-fire silvicultural operations in pine forest stands in the Mediterranean Basin.