Tree-ring records can provide valuable information to advance our understanding of contemporary terrestrial carbon cycling and to reconstruct key metrics in the decades preceding monitoring data. The growing use of tree rings in carbon-cycle research is being facilitated by increasing recognition of reciprocal benefits among research communities. Yet, basic questions persist regarding what tree rings represent at the ecosystem level, how to optimally integrate them with other data streams, and what related challenges need to be overcome. It is also apparent that considerable unexplored potential exists for tree rings to refine assessments of terrestrial carbon cycling across a range of temporal and spatial domains. Here, we summarize recent advances and highlight promising paths of investigation with respect to (1) growth phenology, (2) forest productivity trends and variability, (3) CO2 fertilization and water-use efficiency, (4) forest disturbances, and (5) comparisons between observational and computational forest productivity estimates. We encourage the integration of tree-ring data: with eddy-covariance measurements to investigate carbon allocation patterns and water-use efficiency; with remotely sensed observations to distinguish the timing of cambial growth and leaf phenology; and with forest inventories to develop continuous, annually-resolved and long-term carbon budgets. In addition, we note the potential of tree-ring records and derivatives thereof to help evaluate the performance of earth system models regarding the simulated magnitude and dynamics of forest carbon uptake, and inform these models about growth responses to (non-)climatic drivers. Such efforts are expected to improve our understanding of forest carbon cycling and place current developments into a long-term perspective.

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A tree-ring perspective on the terrestrial carbon cycle

The largest part of forest biomass consists of wood. A global estimate of carbon stored in lignified tissues rises up to 400 Pg. Given these quantities, there is a growing interest of implementing wood research in diagnoses and evaluations of the carrying capacity of the global ecosystem and its forests. The question arises how disciplines like wood anatomy could respond to the increasing demands of a trait-based ecology, understood as a paradigmatic shift in addressing global changes. Dendrochronology and ecological wood anatomy, traditionally operating within the paradigm of species-based ecology, developed robust methods to address ecological questions. However, sampling strategies and database design will likely be different when wood traits are to be used to study individual tree performance, including responses to stress.Aiming at optimally involving wood research in trait-based ecology, some trait concepts are analysed. The value of the IAWA standard lists of wood anatomical features as starting points for trait databases is recognized. A summary of the functionality of wood is given to inform the trait-research community of basic aspects of tree performance. The time dimension is highlighted, as well as the foundations for understanding bio-hydraulics, bio-mechanics and metabolism of wood and relevant traits.Guidelines are given for sampling strategies and database concepts. Prospects of time axis construction and system integration are discussed, as well as the importance of standardizing for size.

Wood anatomy and trait-based ecology

Developmental initiation of plant vascular tissue, including xylem and phloem, from the vascular cambium depends on environmental factors, such as temperature and precipitation. Proper formation of vascular tissue is critical for the transpiration stream, along with photosynthesis as a whole. While effects of individual environmental factors on the transpiration stream are well studied, interactive effects of multiple stress factors are underrepresented. As expected, climate change will result in plants experiencing multiple co-occurring environmental stress factors, which require further studies. Also, the effects of the main climate change components (carbon dioxide, temperature, and drought) on vascular cambium are not well understood. This review aims at synthesizing current knowledge regarding the effects of the main climate change components on the initiation and differentiation of vascular cambium, the transpiration stream, and photosynthesis. We predict that combined environmental factors will result in increased diameter and density of xylem vessels or tracheids in the absence of water stress. However, drought may decrease the density of xylem vessels or tracheids. All interactive combinations are expected to increase vascular cell wall thickness, and therefore increase carbon allocation to these tissues. A comprehensive study of the effects of multiple environmental factors on plant vascular tissue and water regulation should help us understand plant responses to climate change.

https://www.mdpi.com/2223-7747/8/3/65/pdf

Environmental Factors Influence Plant Vascular System and Water Regulation

A better understanding of the influence of environmental conditions on wood formation should help to improve the radial growth of trees and to prepare for climate change.
 The cambial activity of trees is associated with seasonal cycles of activity and dormancy in temperate zones. The timing of cambial reactivation in early spring and dormancy in autumn plays an important role in determination of the cambial growth and the environmental adaptivity of temperate trees. This review focuses on the temperature regulation of the timing of cambial reactivation and xylem differentiation and highlights recent advances of bud growth in relation to cambial activity of temperate trees. In addition, we discuss relationships between the timing of cambial reactivation, start of xylem differentiation and changes in levels of storage materials to identify the source of the energy required for cell division and differentiation. We also present a summary of current understanding of the effects of rapid increases and decreases in temperature on cambial activity, by localized heating and cooling, respectively. Increases in temperature from late winter to early spring influence the physiological processes that are involved in the initiation of cambial reactivation and xylem differentiation both in localized heated stems and under natural conditions. Localized cooling has a direct effect on cell expansion, the thickening of walls of differentiating tracheids, and the rate of division of cambial cells. A rapid decrease in temperature of the stem might be the critical factor in the control of latewood formation and the cessation of cambial activity. Therefore, temperature is the main driver of cambial activity in temperate trees and trees are able to feel changes in temperature through the stem. The climate change might affect wood formation in trees.

Climate change and the regulation of wood formation in trees by temperature

We investigate cambial growth periodicity in Brachystegia spiciformis, a dominant tree species in the seasonally dry miombo woodland of southern Africa. To better understand how the brevi-deciduous (experiencing a short, drought-induced leaf fall period) leaf phenology of this species can be linked to a distinct period of cambial activity, we applied a bi-weekly pinning to six trees in western Zambia over the course of one year. Our results show that the onset and end of cambial growth was synchronous between trees, but was not concurrent with the onset and end of the rainy season. The relatively short (three to four months maximum) cambial growth season corresponded to the core of the rainy season, when 75% of the annual precipitation fell, and to the period when the trees were at full photosynthetic capacity. Tree-ring studies of this species have found a significant relationship between annual tree growth and precipitation, but we did not observe such a correlation at intra-annual resolution in this study. Furthermore, a substantial rainfall event occurring after the end of the cambial growth season did not induce xylem initiation or false ring formation. Low sample replication should be taken into account when interpreting the results of this study, but our findings can be used to refine the carbon allocation component of process-based terrestrial ecosystem models and can thus contribute to a more detailed estimation of the role of the miombo woodland in the terrestrial carbon cycle. Furthermore, we provide a physiological foundation for the use of Brachystegia spiciformis tree-ring records in paleoclimate research.

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Cambial growth season of brevi-deciduous Brachystegia spiciformis trees from south central Africa restricted to less than four months.

s. Birmensdorf: Swiss Federal Research Institute WSL. Hacke UG, Sperry JS, Pockman WT, Davis SD, MFCulloh KA. 2001. Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure. Oecologia 126: 457–461. IAWA (International Association of Wood Anatomists) Committee. 1989. IAWA list of microscopic features for hardwood identification. IAWA Bulletin, New Series 10: 219 – 329. Jacoby GC, D’Arrigo RD. 1990. Teak (Tectona grandis L.f.), a tropical species of large-scale dendroclimatic potential. Dendrochronologia 8:

PART OF A HIGHLIGHT ON TREE BIOLOGY Fluctuations of cambial activity in relation to precipitation result in annual rings and intra-annual growth zones of xylem and phloem in teak (Tectona grandis) in Ivory Coast

Kouame

Papers

PART OF A HIGHLIGHT ON TREE BIOLOGY Fluctuations of cambial activity in relation to precipitation result in annual rings and intra-annual growth zones of xylem and phloem in teak (Tectona grandis) in Ivory Coast