Anthony R. Ambrose

Seasonal and episodic moisture controls on plant and microbial contributions to soil respiration

Moisture inputs drive soil respiration (SR) dynamics in semi-arid and arid ecosystems. However, determining the contributions of root and microbial respiration to SR, and their separate temporal responses to periodic drought and water pulses, remains poorly understood. This study was conducted in a pine forest ecosystem with a Mediterranean-type climate that receives seasonally varying precipitation inputs from both rainfall (in the winter) and fog-drip (primarily in the summer). We used automated SR measurements, radiocarbon SR source partitioning, and a water addition experiment to understand how SR, and its separate root and microbial sources, respond to seasonal and episodic changes in moisture. Seasonal changes in SR were driven by surface soil water content and large changes in root respiration contributions. Superimposed on these seasonal patterns were episodic pulses of precipitation that determined the short-term SR patterns. Warm season precipitation pulses derived from fog-drip, and rainfall following extended dry periods, stimulated the largest SR responses. Microbial respiration dominated these SR responses, increasing within hours, whereas root respiration responded more slowly over days. We conclude that root and microbial respiration sources respond differently in timing and magnitude to both seasonal and episodic moisture inputs. These findings have important implications for the mechanistic representation of SR in models and the response of dry ecosystems to changes in precipitation patterns.

Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood (Sequoia sempervirens) and giant sequoia (Sequoiadendron giganteum) trees of different sizes. Leaf-specific hydraulic conductivity (k(L)) increased with height in S. sempervirens but not in S. giganteum, while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios (delta(13)C) increased, and maximum mass-based stomatal conductance (g(mass)) and photosynthesis (A(mass)) decreased with height in both species. As a result, both A(mass) and g(mass) were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum. In addition, A(mass) and g(mass) were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO(2) conductance (g(i)). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat.

Effects of height on treetop transpiration and stomatal conductance in coast redwood (Sequoia sempervirens)

Treetops become increasingly constrained by gravity-induced water stress as they approach maximum height. Here we examine the effects of height on seasonal and diurnal sap flow dynamics at the tops of 12 unsuppressed Sequoia sempervirens (D. Don) Endl. (coast redwood) trees 68-113 m tall during one growing season. Average treetop sap velocity (V(S)), transpiration per unit leaf area (E(L)) and stomatal conductance per unit leaf area (G(S)) significantly decreased with increasing height. These differences in sap flow were associated with an unexpected decrease in treetop sapwood area-to-leaf area ratios (A(S):A(L)) in the tallest trees. Both E(L) and G(S) declined as soil moisture decreased and vapor pressure deficit (D) increased throughout the growing season with a greater decline in shorter trees. Under high soil moisture and light conditions, reference G(S) (G(Sref); G(S) at D = 1 kPa) and sensitivity of G(S) to D (-δ; dG(S)/dlnD) significantly decreased with increasing height. The close relationship we observed between G(Sref) and -δ is consistent with the role of stomata in regulating E(L) and leaf water potential (Ψ(L)). Our results confirm that increasing tree height reduces gas exchange of treetop foliage and thereby contributes to lower carbon assimilation and height growth rates as S. sempervirens approaches maximum height.

How do tree structure and old age affect growth potential of California redwoods

As the only species exceeding 90 m in height and 2000 years of age, Sequoia sempervirens and Sequoiadendron giganteum provide the optimal platform upon which to examine interactions among tree structure, age, and growth. We climbed 140 trees in old-growth redwood forests across California, USA, spanning a broad range of sizes and including the tallest, largest, and oldest known living individuals (i.e., 115.86 vs. 96.29 m tall, 424 vs. 582 Mg aboveground dry mass, and 2510 vs. 3240 years old for Sequoia and Sequoiadendron, respectively). We used a combination of direct measurements, hierarchical sampling, and dendrochronology to quantify tree structure and annual growth increments through old age. We also developed equations to predict aboveground attributes of standing redwoods via ground-based measurements. Compared to Sequoia, Sequoiadendron develops thicker bark on lower trunks, provisions leaves with more sapwood, and delays heartwood production throughout the crown. Main trunk wood volume growth (up to 1.6 vs. 0.9 m3/yr), aboveground biomass growth (up to 0.77 vs. 0.45 Mg/yr), and aboveground growth efficiency (0.55 ± 0.04 vs. 0.22 ± 0.01 kg annual growth per kg leaves, mean ± SE) are all higher in Sequoia. Two independent dimensions of structure—size and aboveground vigor—are the strongest predictors of tree-level productivity in both species. A third dimension, relative trunk size, is a significant predictor of growth in Sequoia such that trees with relatively large main trunks compared to their crowns produce more wood annually. Similar-size trees grow at similar rates regardless of latitude or elevation in tall forests of each species. Recent annual growth increments are higher than in the past for the majority of trees, and old trees are just as responsive to environmental changes as young trees. Negative growth–age relationships in previous centuries and positive growth–age relationships in recent decades reflect sampling bias and shifting disturbance regimes. Overall, we find little (if any) evidence for negative effects of old age on tree-level productivity in either species. Except for recovery periods following temporary reductions in crown size, annual increments of wood volume and biomass growth increase as redwoods enlarge with age until extrinsic forces cause tree death.

Cloud shading and fog drip influence the metabolism of a coastal pine ecosystem

Assessing the ecological importance of clouds has substantial implications for our basic understanding of ecosystems and for predicting how they will respond to a changing climate. This study was conducted in a coastal Bishop pine forest ecosystem that experiences regular cycles of stratus cloud cover and inundation in summer. Our objective was to understand how these clouds impact ecosystem metabolism by contrasting two sites along a gradient of summer stratus cover. The site that was under cloud cover ~15% more of the summer daytime hours had lower air temperatures and evaporation rates, higher soil moisture content, and received more frequent fog drip inputs than the site with less cloud cover. These cloud-driven differences in environmental conditions translated into large differences in plant and microbial activity. Pine trees at the site with greater cloud cover exhibited less water stress in summer, larger basal area growth, and greater rates of sap velocity. The difference in basal area growth between the two sites was largely due to summer growth. Microbial metabolism was highly responsive to fog drip, illustrated by an observed ~3-fold increase in microbial biomass C with increasing summer fog drip. In addition, the site with more cloud cover had greater total soil respiration and a larger fractional contribution from heterotrophic sources. We conclude that clouds are important to the ecological functioning of these coastal forests, providing summer shading and cooling that relieve pine and microbial drought stress as well as regular moisture inputs that elevate plant and microbial metabolism. These findings are important for understanding how these and other seasonally dry coastal ecosystems will respond to predicted changes in stratus cover, rainfall, and temperature.

Contrasting drought-response strategies in California redwoods

We compared the physiology and growth of seedlings originating from different Sequoia sempervirens (D. Don.) Endl. (coast redwood) and Sequoiadendron giganteum (Lindl.) Buchh. (giant sequoia) populations subjected to progressive drought followed by a recovery period in a controlled greenhouse experiment. Our objective was to examine how multiple plant traits interact to influence the response of seedlings of each species and seed population to a single drought and recovery cycle. We measured soil and plant water status, leaf gas exchange, stem embolism and growth of control (well-watered) and drought-stressed (water withheld) seedlings from each population at the beginning, middle and end of a 6-week drought period and again 2 weeks after re-watering. The drought had a significant effect on many aspects of seedling performance, but water-stressed seedlings regained most physiological functioning by the end of the recovery period. Sequoiadendron seedlings exhibited a greater degree of isohydry (water status regulation), lower levels of stem embolism, higher biomass allocation to roots and lower sensitivity of growth to drought compared with Sequoia. Only minor intra-specific differences were observed among populations. Our results show that seedlings of the two redwood species exhibit contrasting drought-response strategies that align with the environmental conditions these trees experience in their native habitats, and demonstrate trade-offs and coordination among traits affecting plant water use, carbon gain and growth under drought.