Duncan D. Smith

The roles of hydraulic and carbon stress in a widespread climate-induced forest die-off

Forest ecosystems store approximately 45% of the carbon found in terrestrial ecosystems, but they are sensitive to climate-induced dieback. Forest die-off constitutes a large uncertainty in projections of climate impacts on terrestrial ecosystems, climate–ecosystem interactions, and carbon-cycle feedbacks. Current understanding of the physiological mechanisms mediating climate-induced forest mortality limits the ability to model or project these threshold events. We report here a direct and in situ study of the mechanisms underlying recent widespread and climate-induced trembling aspen (Populus tremuloides) forest mortality in western North America. We find substantial evidence of hydraulic failure of roots and branches linked to landscape patterns of canopy and root mortality in this species. On the contrary, we find no evidence that drought stress led to depletion of carbohydrate reserves. Our results illuminate proximate mechanisms underpinning recent aspen forest mortality and provide guidance for understanding and projecting forest die-offs under climate change.

Hydraulic trade-offs and space filling enable better predictions of vascular structure and function in plants

Plant vascular networks are central to botanical form, function, and diversity. Here, we develop a theory for plant network scaling that is based on optimal space filling by the vascular system along with trade-offs between hydraulic safety and efficiency. Including these evolutionary drivers leads to predictions for sap flow, the taper of the radii of xylem conduits from trunk to terminal twig, and how the frequency of xylem conduits varies with conduit radius. To test our predictions, we use comprehensive empirical measurements of maple, oak, and pine trees and complementary literature data that we obtained for a wide range of tree species. This robust intra- and interspecific assessment of our botanical network model indicates that the central tendency of observed scaling properties supports our predictions much better than the West, Brown, and Enquist (WBE) or pipe models. Consequently, our model is a more accurate description of vascular architecture than what is given by existing network models and should be used as a baseline to understand and to predict the scaling of individual plants to whole forests. In addition, our model is flexible enough to allow the quantification of species variation around rules for network design. These results suggest that the evolutionary drivers that we propose have been fundamental in determining how physiological processes scale within and across plant species.

Rare pits, large vessels and extreme vulnerability to cavitation in a ring-porous tree species

• The rare pit hypothesis predicts that the extensive inter-vessel pitting in large early-wood vessels of ring-porous trees should render many of these vessels extremely vulnerable to cavitation by air-seeding. This prediction was tested in Quercus gambelii. • Cavitation was assessed from native hydraulic conductivity at field sap tension and in dehydrated branches. Single-vessel air injections gave air-seeding pressures through vessel files; these data were used to estimate air-seeding pressures for inter-vessel walls and pits. • Extensive cavitation occurred at xylem sap tensions below 1 MPa. Refilling occurred below 0.5 MPa and was inhibited by phloem girdling. Remaining vessels cavitated over a wide range to above 4 MPa. Similarly, 40% of injected vessel files air-seeded below 1.0 MPa, whereas the remainder seeded over a wide range exceeding 5 MPa. Inter-vessel walls averaged 1.02 MPa air-seeding pressure, similar and opposite to the mean cavitation tension of 1.22 MPa. Consistent with the rare pit hypothesis, only 7% of inter-vessel pits were estimated to air-seed by 1.22 MPa. • The results confirm the rare pit prediction that a significant fraction of large vessels in Q. gambelii experience high probability of failure by air-seeding.

An empirical assessment of tree branching networks and implications for plant allometric scaling models

Several theories predict whole-tree function on the basis of allometric scaling relationships assumed to emerge from traits of branching networks. To test this key assumption, and more generally, to explore patterns of external architecture within and across trees, we measure branch traits (radii/lengths) and calculate scaling exponents from five functionally divergent species. Consistent with leading theories, including metabolic scaling theory, branching is area preserving and statistically self-similar within trees. However, differences among scaling exponents calculated at node- and whole-tree levels challenge the assumption of an optimised, symmetrically branching tree. Furthermore, scaling exponents estimated for branch length change across branching orders, and exponents for scaling metabolic rate with plant size (or number of terminal tips) significantly differ from theoretical predictions. These findings, along with variability in the scaling of branch radii being less than for branch lengths, suggest extending current scaling theories to include asymmetrical branching and differential selective pressures in plant architectures.

Coordination between water transport capacity, biomass growth, metabolic scaling and species stature in co-occurring shrub and tree species

The significance of xylem function and metabolic scaling theory begins from the idea that water transport is strongly coupled to growth rate. At the same time, coordination of water transport and growth seemingly should differ between plant functional types. We evaluated the relationships between water transport, growth and species stature in six species of co-occurring trees and shrubs. Within species, a strong proportionality between plant hydraulic conductance (K), sap flow (Q) and shoot biomass growth (G) was generally supported. Across species, however, trees grew more for a given K or Q than shrubs, indicating greater growth-based water-use efficiency (WUE) in trees. Trees also showed slower decline in relative growth rate (RGR) than shrubs, equivalent to a steeper G by mass (M) scaling exponent in trees (0.77-0.98). The K and Q by M scaling exponents were common across all species (0.80, 0.82), suggesting that the steeper G scaling in trees reflects a size-dependent increase in their growth-based WUE. The common K and Q by M exponents were statistically consistent with the 0.75 of ideal scaling theory. A model based upon xylem anatomy and branching architecture consistently predicted the observed K by M scaling exponents but only when deviations from ideal symmetric branching were incorporated.

A synthesis of the effects of atmospheric carbon dioxide enrichment on plant hydraulics: implications for whole-plant water use efficiency and resistance to drought†

Here, we summarize studies on the effects of elevated [CO2 ] (CO2e ) on the structure and function of plant hydraulic architecture and explore the implications of those changes using a model. Changes in conduit diameter and hydraulic conductance due to CO2e vary among species. Ring-porous species tend towards an increase in conduit size and consequently conductivity. The effect in diffuse-porous species is much more limited. In conifers, the results are mixed, some species show minor changes in xylem structure, while other studies found increases in tracheid density and diameter. Non-woody plants generally exhibited the reverse pattern with narrower conduits and lower hydraulic conductivity under CO2e . Further, changes in drought-resistance traits suggest that non-woody plants were the most affected by CO2e , which may permit them to better resist drought-induced embolism under future conditions. Due to their complexity, acclimation in hydraulic traits in response to CO2e is difficult to interpret when relying solely on measurements. When we examined how the observed tissues-specific trends might alter plant function, our modelling results suggest that these hydraulic changes would lead to reduced conductance and more frequent drought stress in trees that develop under CO2e with a more pronounced effect in isohydric than in anisohydric species.

Convergence in leaf size versus twig leaf area scaling: do plants optimize leaf area partitioning?

Background and Aims Corner’s rule states that thicker twigs bear larger leaves. The exact nature of this relationship and why it should occur has been the subject of numerous studies. It is obvious that thicker twigs should support greater total leaf area () for hydraulical and mechanical reasons. But it is not obvious why mean leaf size () should scale positively with . We asked what this scaling relationship is within species and how variable it is across species. We then developed a model to explain why these relationships exist. Methods To minimize potential sources of variability, we compared twig properties from six co-occurring and functionally similar species: Acer grandidentatum, Amelanchier alnifolia, Betula occidentalis, Cornus sericea, Populus fremontii and Symphoricarpos oreophilus. We modelled the economics of leaf display, weighing the benefit from light absorption against the cost of leaf tissue, to predict the optimal combinations under different canopy openings. Key Results We observed a common by exponent of 0.6, meaning that and leaf number on twigs increased in a specific coordination. Common scaling exponents were not supported for relationships between any other measured twig properties. The model consistently predicted positive by scaling when twigs optimally filled canopy openings. The observed 0·6 exponent was predicted when self-shading decreased with larger canopy opening. Conclusions Our results suggest Corner’s rule may be better understood when recast as positive by scaling. Our model provides a tentative explanation of observed by scaling and suggests different scaling may exist in different environments.