Heidi J. Renninger

Physiological strategies of co-occurring oaks in a water- and nutrient-limited ecosystem.

Oak species are well suited to water-limited conditions by either avoiding water stress through deep rooting or tolerating water stress through tight stomatal control. In co-occurring species where resources are limited, species may either partition resources in space and/or time or exhibit differing efficiencies in the use of limited resources. Therefore, this study seeks to determine whether two co-occurring oak species (Quercus prinus L. and Quercus velutina Lam.) differ in physiological parameters including photosynthesis, stomatal conductance, water-use (WUE) and nitrogen-use efficiency (NUE), as well as to characterize transpiration and average canopy stomatal responses to climatic variables in a sandy, well-drained and nutrient-limited ecosystem. The study was conducted in the New Jersey Pinelands and we measured sap flux over a 3-year period, as well as leaf gas exchange, leaf nitrogen and carbon isotope concentrations. Both oak species showed relatively steep increases in leaf-specific transpiration at low vapor pressure deficit (VPD) values before maximum transpiration rates were achieved, which were sustained over a broad range in VPD. This suggests tight stomatal control over transpiration in both species, although Q. velutina showed significantly higher leaf-level and canopy-level stomatal conductance than Q. prinus. Average daytime stomatal conductance was positively correlated with soil moisture and both oak species maintained at least 75% of their maximum canopy stomatal conductance at soil moistures in the upper soil layer (0-0.3 m) as low as 0.03 m(3) m(3)(-3). Quercus velutina had significantly higher photosynthetic rates, maximum Rubisco-limited and electron-transport-limited carboxylation rates, dark respiration rates and nitrogen concentration per unit leaf area than Q. prinus. However, both species exhibited similar WUEs and NUEs. Therefore, Q. prinus has a more conservative resource-use strategy, while Q. velutina may need to exploit niches that are locally higher in nutrients and water. Likewise, both species appear to tap deep, stable water sources, highlighting the importance of rooting depth in modeling transpiration and stomatal conductance in many oak ecosystems.

Comparison of Tissue Heat Balance- and Thermal Dissipation-Derived Sap Flow Measurements in Ring-Porous Oaks and a Pine

Sap flow measurements have become integral in many physiological and ecological investigations. A number of methods are used to estimate sap flow rates in trees, but probably the most popular is the thermal dissipation (TD) method because of its affordability, relatively low power consumption, and ease of use. However, there have been questions about the use of this method in ring-porous species and whether individual species and site calibrations are needed. We made concurrent measurements of sap flow rates using TD sensors and the tissue heat balance (THB) method in two oak species (Quercus prinus Willd. and Quercus velutina Lam.) and one pine (Pinus echinata Mill.). We also made concurrent measurements of sap flow rates using both 1 and 2-cm long TD sensors in both oak species. We found that both the TD and THB systems tended to match well in the pine individual, but sap flow rates were underestimated by 2-cm long TD sensors in five individuals of the two ring-porous oak species. Underestimations of 20–35% occurred in Q. prinus even when a “Clearwater” correction was applied to account for the shallowness of the sapwood depth relative to the sensor length and flow rates were underestimated by up to 50% in Q. velutina. Two centimeter long TD sensors also underestimated flow rates compared with 1-cm long sensors in Q. prinus, but only at large flow rates. When 2-cm long sensor data in Q. prinus were scaled using the regression with 1-cm long data, daily flow rates matched well with the rates measured by the THB system. Daily plot level transpiration estimated using TD sap flow rates and scaled 1 cm sensor data averaged about 15% lower than those estimated by the THB method. Therefore, these results suggest that 1-cm long sensors are appropriate in species with shallow sapwood, however more corrections may be necessary in ring-porous species.

Comparative hydraulic and anatomic properties in palm trees (Washingtonia robusta) of varying heights: implications for hydraulic limitation to increased height growth

As trees grow taller, the energetic cost of moving water to the leaves becomes higher and could begin to limit carbon gain and subsequent growth. The hydraulic limitation hypothesis states that as trees grow taller, the path length and therefore frictional resistance of water flow increases, leading to stomatal closure, reduced photosynthesis and decreased height growth in tall trees. Although this hypothesis is supported by the physical laws governing water movement in trees, its validation has been complicated by the complex structure of most tree species. Therefore, this study tested the hydraulic limitation hypothesis in Washingtonia robusta (H. Wendl.), a palm that, while growing to tall heights, is still structurally simple enough to act as a model organism for testing. There were no discernable relationships between tree height and stomatal conductance, stomatal densities, guard cell lengths, leaf dry mass per unit area (LMA) or sap flux, suggesting that these key aspects of hydraulic limitation are not reduced in taller palms. Taller palms did, however, have higher maximum daily photosynthetic assimilation rates, lower minimum leaf water potentials that occurred earlier in the day and fewer, smaller leaves than did shorter palms. Leaf epidermal cells were also smaller in taller palms compared with shorter ones. These findings are consistent with hydraulic compensation in that tall palms may be overcoming the increased path length resistance through smaller, more efficient leaves and lower leaf water potentials than shorter palms.

Resource use and efficiency, and stomatal responses to environmental drivers of oak and pine species in an Atlantic Coastal Plain forest

Pine-oak ecosystems are globally distributed even though differences in anatomy and leaf habit between many co-occurring oaks and pines suggest different strategies for resource use, efficiency and stomatal behavior. The New Jersey Pinelands contain sandy soils with low water- and nutrient-holding capacity providing an opportunity to examine trade-offs in resource uptake and efficiency. Therefore, we compared resource use in terms of transpiration rates and leaf nitrogen content and resource-use efficiency including water-use efficiency (WUE) via gas exchange and leaf carbon isotopes and photosynthetic nitrogen-use efficiency (PNUE) between oaks (Quercus alba, Q. prinus, Q. velutina) and pines (Pinus rigida, P. echinata). We also determined environmental drivers [vapor pressure deficit (VPD), soil moisture, solar radiation] of canopy stomatal conductance (GS) estimated via sap flow and stomatal sensitivity to light and soil moisture. Net assimilation rates were similar between genera, but oak leaves used about 10% more water and pine foliage contained about 20% more N per unit leaf area. Therefore, oaks exhibited greater PNUE while pines had higher WUE based on gas exchange, although WUE from carbon isotopes was not significantly different. For the environmental drivers of GS, oaks had about 10% lower stomatal sensitivity to VPD normalized by reference stomatal conductance compared with pines. Pines exhibited a significant positive relationship between shallow soil moisture and GS, but only GS in Q. velutina was positively related to soil moisture. In contrast, stomatal sensitivity to VPD was significantly related to solar radiation in all oak species but only pines at one site. Therefore, oaks rely more heavily on groundwater resources but have lower WUE, while pines have larger leaf areas and nitrogen acquisition but lower PNUE demonstrating a trade-off between using water and nitrogen efficiently in a resource-limited ecosystem.

Modeling respiration from snags and coarse woody debris before and after an invasive gypsy moth disturbance

Although snags and coarse woody debris are a small component of ecosystem respiration, disturbances can significantly increase the mass and respiration from these carbon (C) pools. The objectives of this study were to (1) measure respiration rates of snags and coarse woody debris throughout the year in a forest previously defoliated by gypsy moths, (2) develop models for dead stem respiration rates, (3) model stand-level respiration rates of dead stems using forest inventory and analysis data sets and environmental variables predisturbance and postdisturbance, and (4) compare total dead stem respiration rates with total ecosystem respiration and net ecosystem exchange. Respiration rates were measured on selected Pinus and Quercus snags and coarse woody debris each month for 1 year in a northeastern U.S. temperate forest. Multiple linear regression using environmental and biometric variables including wood temperature, diameter, density, species, and decay class was used to model respiration rates of dead stems. The mass of snags and coarse woody debris increased more than fivefold after disturbance and respiration rates increased more than threefold. The contribution of dead stems to total ecosystem respiration more than tripled from 0.85% to almost 3% and respiration from dead stems alone was approximately equal to the net ecosystem exchange of the forest in 2011 (fourth year postdisturbance). This study highlights the importance of dead stem C pools and fluxes particularly during disturbance and recovery cycles. With climate change increasing the ranges of many forest pests and pathogens, these data become particularly important for accurately modeling future C cycling.

NO CORRELATION BETWEEN LATEWOOD FORMATION AND LEADER GROWTH IN DOUGLAS-FIR SAPLINGS

SUMMARY The width of earlywood and latewood in conifer xylem may have a profound effect on water transport and storage, vulnerability to embolism, and wood strength, yet the controls over the timing of latewood formation are unclear. Tracheids differentiating in the cambial zone are influenced by IAA indole-3 acetic acid, the radial concentration gradient of which appears to either increase cell expansion (earlywood) or increase cell wall deposition (latewood). There are suggestive data that latewood begins to form when the growth of the leader stops, but defini tive results are lacking. Height growth was measured in 14 Douglas-fir (Pseudotsuga menziesii) saplings at 10 dates between May and August, from the beginning of the growing season until after height growth had ceased. The cambium was also pinned six times between June and July, to induce xylem scarring at known dates. After height growth ceased, saplings were harvested and transverse sections of the wood were made at the pin insertion points. The date at which 95% of the height growth had occurred and the date at which latewood formation had begun were estimated. Analysis showed no correlation of these data, suggesting that the two phenomena may occur around the same time, but that one is not causal of the other.

Hydraulic properties of fronds from palms of varying height and habitat

Because palms grow in highly varying climates and reach considerable heights, they present a unique opportunity to evaluate how environment and plant size impact hydraulic function. We studied hydraulic properties of petioles from palms of varying height from three species: Iriartea deltoidea, a tropical rainforest species; Mauritia flexuosa, a tropical rainforest, swamp species; and Washingtonia robusta, a subtropical species. We measured leaf areas, petiole cross-sectional areas, specific conductivity (KS), petiole anatomical properties, vulnerability to embolism and leaf water potentials and calculated petiole Huber values and leaf-specific conductivities (KL). Leaf and petiole cross-sectional areas varied widely with height. However, hydraulic properties including Huber values, KS and KL, remained constant. The two palmate species, M. flexuosa and W. robusta, had larger Huber values than I. deltoidea, a pinnately-compound species which exhibited the highest KS. Metaxylem vessel diameters and vascular bundle densities varied with height in opposing patterns to maintain petiole conductivities. I. deltoidea and W. robusta petioles had similar P50 values (the point at which 50% of hydraulic conductivity is lost) averaged over all crown heights, but W. robusta exhibited more negative P50 values in taller palms. Comparison of P50 values with transpiring midday leaf water potentials, as well as a double-dye staining experiment in a 1-m-tall palm, suggested that a fairly significant amount of embolisms were occurring and refilled on a diurnal basis. Therefore, across palms differing widely in height and growing environments, we found convergence in water transport per unit leaf area (KL) with individuals exhibiting differing strategies for achieving this.

Intrinsic and extrinsic hydraulic factors in varying sizes of two Amazonian palm species (Iriartea deltoidea and Mauritia flexuosa) differing in development and growing environment.

UNLABELLED PREMISE OF THE STUDY This study seeks to determine how hydraulic factors vary with ontogeny and whether they begin to limit further height growth in palms. Palms are an attractive group for physiological research because their columnar trunks and simple leaf habit allow key intrinsic and extrinsic hydraulic variables to be estimated more easily than in complex arborescent dicotyledons. • METHODS We measured various biometric and physiological factors including sap flux, leaf areas, turnover rates, and internode lengths in two Amazonian rainforest species: terra firme Iriartea deltoidea and swamp-adapted Mauritia flexuosa. These two palm species differ markedly in edaphic conditions, leaf type (pinnately compound vs. palmate), and bole development, making physiological comparisons between them important as well. • KEY RESULTS The species exhibited differing patterns in height growth rate along boles, which appear to relate to their differences in bole development. Growth rates ultimately slowed at the tops of tall palms in both species. We also found a high degree of convergence in total leaf area with height in both species even though they exhibited contrasting patterns in both live frond number and leaf area per frond with height. Sap flux density from leaves was constant with height but four times greater in M. flexuosa than in I. deltoidea. • CONCLUSIONS Although height growth rates slow considerably in tall palms, neither species shows evidence that hydraulic factors become limiting because they are able to support much greater leaf areas with similar sap flux densities as shorter palms.

A comparison of the hydraulic efficiency of a palm species (Iriartea deltoidea) with other wood types

Palms are an important component of tropical ecosystems, living alongside dicotyledonous trees, even though they have a very different growth pattern and vascular system. As monocots, vessels in palms are located within vascular bundles and, without a vascular cambium that many dicotyledonous trees possess, palms cannot add additional vessels to their vascular system as they get older and taller. This means that hydraulic architecture in palms is more predetermined, which may require a highly efficient hydraulic system. This preset nature, along with the decoupling of hydraulic and mechanical functioning to different cell types, may allow palms to have a more efficient hydraulic system than dicotyledonous trees. Therefore, this study seeks to determine the efficiency of the hydraulic system in the palm Iriartea deltoidea (Ruiz & Pav.) and compare this efficiency with other tree forms. We measured cross-sectional areas of roots, stems and fronds as well as leaf areas of I. deltoidea saplings. Likewise, cross-sections were made and vessel diameters and frequencies measured. This allowed for the calculation of theoretical specific conductivity (K(S,calc)), theoretical leaf-specific conductivity (K(L,calc)), and vessel diameter and vessel number ratios between distal and proximal locations in the palms. Iriartea deltoidea palms were found to have the largest, least frequent vessels that diverged most from the square packing limit (maximum number of vessels that fit into a given area) compared with other major tree forms, and they therefore invested the least space and carbon into water transport structures. Likewise, conduits tapered by ∼1/3 between ranks (root, bole and petiole), which represents an efficient ratio with regard to the trade-offs between safety and efficiency of the conducting system. Conduits also exhibited a high conservation of the sum of the conduit radii cubed (Σr(3)) across ranks, thereby approximating Murrays law patterning. Therefore, our results indicate that the palm I. deltoidea has a very efficient hydraulic system in terms of maintaining a large conducting capacity with a minimal vascular investment. This efficiency may allow palms to compete well with dicotyledonous trees in tropical and subtropical climates but other developmental factors largely restrict palms from regions that experience prolonged freezing temperatures.

Forest response and recovery following disturbance in upland forests of the Atlantic Coastal Plain

Carbon and water cycling of forests contribute significantly to the Earths overall biogeochemical cycling and may be affected by disturbance and climate change. As a larger body of research becomes available about leaf-level, ecosystem and regional scale effects of disturbances on forest ecosystems, a more mechanistic understanding is developing which can improve modeling efforts. Here, we summarize some of the major effects of physical and biogenic disturbances, such as drought, prescribed fire, and insect defoliation, on leaf and ecosystem-scale physiological responses as well as impacts on carbon and water cycling in an Atlantic Coastal Plain upland oak/pine and upland pine forest. During drought, stomatal conductance and canopy stomatal conductance were reduced, however, defoliation increased conductance on both leaf-level and canopy scale. Furthermore, after prescribed fire, leaf-level stomatal conductance was unchanged for pines but decreased for oaks, while canopy stomatal conductance decreased temporarily, but then rebounded the following growing season, thus exhibiting transient responses. This study suggests that forest response to disturbance varies from the leaf to ecosystem level as well as species level and thus, these differential responses interplay to determine the fate of forest structure and functioning post disturbance.