Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θv), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in θv was observed at 90–120 cm depth in both years. Mean soil water δD during the dry season was −19‰ at 0–30 cm, −34‰ at 30–60 cm, and −50‰ at 90–120 cm. Average values of xylem δD among the liana species ranged from –28‰ to –44‰ during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem δD values (–41‰ to –62‰), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size

Height-related trends in leaf xylem anatomy and shoot hydraulic characteristics in a tall conifer: safety versus efficiency in foliar water transport

• Hydraulic vulnerability of Douglas-fir (Pseudotsuga menziesii) branchlets decreases with height, allowing shoots at greater height to maintain hydraulic conductance (Kshoot) at more negative leaf water potentials (Ψl). • To determine the basis for this trend shoot hydraulic and tracheid anatomical properties of foliage from the tops of Douglas-fir trees were analysed along a height gradient from 5 to 55 m. • Values of Ψl at which Kshoot was substantially reduced, declined with height by 0.012 Mpa m−1. Maximum Kshoot was reduced by 0.082 mmol m−2 MPa−1 s−1 for every 1 m increase in height. Total tracheid lumen area per needle cross-section, hydraulic mean diameter of leaf tracheid lumens, total number of tracheids per needle cross-section and leaf tracheid length decreased with height by 18.4 μm2 m−1, 0.029 μm m−1, 0.42 m−1 and 5.3 μm m−1, respectively. Tracheid thickness-to-span ratio (tw/b)2 increased with height by 1.04 × 10–3 m−1 and pit number per tracheid decreased with height by 0.07 m−1. • Leaf anatomical adjustments that enhanced the ability to cope with vertical gradients of increasing xylem tension were attained at the expense of reduced water transport capacity and efficiency, possibly contributing to height-related decline in growth of Douglas fir.Keywords: growth limitation, hydraulic conductance, water stress, foliar anatomy, Pseudotsuga menziesii, embolismKeywords: growth limitation, hydraulic conductance, water stress, foliar anatomy, Pseudotsuga menziesii, embolis

Variações em atributos biofísicos e fisiológicos em nível de individuo e de parcela ao longo de um gradiente de densidade arbórea no Cerrado

O objetivo deste estudo foi caracterizar e compreender as variações morfológicas e fisiológicas em atributos relacionados à economia da água, em nível de indivíduo e de parcela, que ocorrem ao longo de um gradiente de densidade arbórea em uma savana neotropical (cerrado). Parcelas de vegetação de cerrado submetidas a um experimento de adubação de longo prazo e matas de galeria também foram incluídas na análise, a fim de expandir os eixos de variação do ecossistema. Mudanças consistentes em atributos biofísicos em nível de parcela foram observadas ao longo do gradiente de densidade crescente de árvores. Estas incluíram uma diminuição na densidade média ponderada da madeira, aumentos na área foliar por planta, na condutividade hidráulica específica da folha, na área foliar específica e na condutância estomática. Um modelo conceitual das interações entre as características biofísicas, morfológicas e fisiológicas foi desenvolvido em uma tentativa de explicar os determinantes da arquitetura hidráulica e variações na economia de água de árvores do Cerrado. Consistente com o modelo, os valores mínimos de potencial hídrico durante a estação seca foram maiores nas árvores de baixa densidade de madeira, em comparação com árvores de alta densidade de madeira, enquanto os maiores valores de potencial hídrico foliar estiveram associados com maior condutância estomática. Por outro lado, ao longo do gradiente de densidade arbórea, as variações em nível de parcela das características morfológicas e fisiológicas não puderam ser explicadas por variações de um único fator ambiental. Alguns dos fatores contribuintes potenciais são: maior concentração e disponibilidade de nutrientes do solo na porção superior do gradiente de densidade arbórea onde há um maior número de árvores e restrições no estabelecimento e crescimento pela inundação periódica e pela presença de concreções no perfil do solo onde a densidade de árvores é mais baixa.The objective of this study was to characterize and understand morphological and physiological variation in traits related to water economy, at the plant and stand level, along a gradient of tree density in a Neotropical savanna (Cerrado). Cerrado plots subjected to long-term fertilization and gallery forests were also included in the analysis to expand the axes of ecosystem variation. Consistent changes in stand level biophysical traits observed along the gradient of increasing tree density included a decrease in weighted-average wood density, and increases in leaf surface area per plant, leaf specific hydraulic conductivity, specific leaf area and stomatal conductance. A conceptual model of biophysical, morphological and physiological trait interactions was developed in an attempt to explain determinants of hydraulic architecture and variations in water economy of Cerrado trees. Consistent with the model minimum leaf water potentials were higher, during the dry season, in low wood density trees compared to high wood density trees, and higher leaf water potential was associated with higher stomatal conductance. On the other hand, variations in stand level physiological and morphological traits along the tree density gradient could not be explained by variations in single environmental factors. Some of the potential contributing factors are: higher concentration and availability of soil nutrients in the upper portion of the gradient where tree density is greatest and constrains to tree establishment and growth by waterlogging and presence of concretions in the soil profile where tree density is lowest

Relações hídricas e arquitetura hidráulica em árvores do cerrado : adequação às variações sazonais de disponibilidade hídrica e de demanda evaporativa

O objetivo deste estudo foi determinar os ajustamentos na morfologia e fisiologia que permitem árvores das savanas neotropicais do Brasil Central (Cerrado) de evitar déficits hídricos e de manter um balanço hídrico interno praticamente constante apesar das variações sazonais da precipitação e no déficit de saturação do ar (D). A precipitação na área de estudo é fortemente sazonal, com cerca de cinco meses praticamente sem chuva durante os quais D é duas vezes maior aos valores medidos na época chuvosa. Como conseqüência da flutuação sazonal das chuvas e de D, o potencial hídrico do solo muda substancialmente, nos primeiros 100 cm do solo, mas permanece quase constante abaixo de 2 m de profundidade. A arquitetura hidráulica e os parâmetros relacionados a relações hídricas das árvores do Cerrado se ajustaram durante a estação seca para evitar o déficit hídrico crescente e assegurar a homeostase nos valores mínimos de potencial hídrico foliar ΨL e na perda total diária de água pela planta (iso-hidria). O comportamento iso-hídrico das árvores do Cerrado foi o resultado de uma diminuição da superfície foliar total por árvore, um forte controle estomático das perdas por evaporação, um aumento na condutividade hidráulica específica da folha e na condutância hidráulica foliar e um aumento da quantidade de águas retirada dos reservatórios internos do caule, durante a estação seca. A eficiência no transporte de água aumentou, nas mesmas proporções, nas folhas e nos ramos terminais durante a estação seca. Todos estes ajustamentos sazonais foram importantes para a manutenção de ΨL acima de limiares críticos, com isto contribuindo para uma redução na formação de embolismos nos ramos e ajudando a evitar a perda de turgor em tecidos foliares durante a época seca. Esses ajustes permitem que os ramos das espécies lenhosas do Cerrado operem bem distanciados do ponto de disfunção catastrófica para a cavitação, enquanto as folhas operam próximas e sofrem embolismos em uma base diária, especialmente durante a estação seca.We determined adjustments in physiology and morphology that allow Neotropical savanna trees from central Brazil (Cerrado) to avoid water deficits and to maintain a nearly constant internal water balance despite seasonal changes in precipitation and air saturation deficit (D). Precipitation in the study area is highly seasonal with about five nearly rainless months during which D is two fold higher compared to wet season values. As a consequence of the seasonal fluctuations in rainfall and D, soil water potential changes substantially in the upper 100 cm of soil, but remains nearly constant below 2 m depth. Hydraulic architecture and water relations traits of Cerrado trees adjusted during the dry season to prevent increasing water deficits and insure homeostasis in minimum leaf water potential ΨL and in total daily water loss per plant (isohydry). The isohydric behavior of Cerrado trees was the result of a decrease in total leaf surface area per tree, a strong stomatal control of evaporative losses, an increase in leaf-specific hydraulic conductivity and leaf hydraulic conductance and an increase in the amount of water withdrawn from internal stem storage, during the dry season. Water transport efficiency increased in the same proportion in leaves and terminal stems during the dry season. All of these seasonal adjustments were important for maintaining ΨL above critical thresholds, which reduces the rate of embolism formation in stems and help to avoid turgor loss in leaf tissues still during the dry season. These adjustments allow the stems of most Cerrado woody species to operate far from the point of catastrophic dysfunction for cavitation, while leaves operate close to it and experience embolism on a daily basis, especially during the dry season

Reduced wood stiffness and strength, and altered stem form, in young antisense 4CL transgenic poplars with reduced lignin contents

• Reduced lignin content in perennial crops has been sought as a means to improve biomass processability for paper and biofuels production, but it is unclear how this could affect wood properties and tree form. • Here, we studied a nontransgenic control and 14 transgenic events containing an antisense 4-coumarate:coenzyme A ligase (4CL) to discern the consequences of lignin reduction in poplar (Populus sp.). During the second year of growth, trees were grown either free-standing in a field trial or affixed to stakes in a glasshouse. • Reductions in lignin of up to 40% gave comparable losses in wood strength and stiffness. This occurred despite the fact that low-lignin trees had a similar wood density and up to three-fold more tension wood. In free-standing and staked trees, the control line had twice the height for a given diameter as did low-lignin trees. Staked trees had twice the height for a given diameter as free-standing trees in the field, but did not differ in wood stiffness. • Variation in tree morphogenesis appears to be governed by lignin · environment interactions mediated by stresses exerted on developing cells. Therefore our results underline the importance of field studies for assessing the performance of transgenic trees with modified wood properties.Keywords: tension wood, lignin, stem form, wood strength, buckling safety factor, transgenic poplar, wood stiffnes

Coordination of leaf structure and gas exchange along a height gradient in a tall conifer

The gravitational component of water potential and frictional resistance during transpiration lead to substantial reductions in leaf water potential (Ψl) near the tops of tall trees, which can influence both leaf growth and physiology. We examined the relationships between morphological features and gas exchange in foliage collected near the tops of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees of different height classes ranging from 5 to 55 m. This sampling allowed us to investigate the effects of tree height on leaf structural characteristics in the absence of potentially confounding factors such as irradiance, temperature, relative humidity and branch length. The use of cut foliage for measurement of intrinsic gas-exchange characteristics allowed identification of height-related trends without the immediate influences of path length and gravity. Stomatal density, needle length, needle width and needle area declined with increasing tree height by 0.70 mm−2 m−1, 0.20 mm m−1, 5.9 × 10−3 mm m−1 and 0.012 mm2 m−1, respectively. Needle thickness and mesophyll thickness increased with tree height by 4.8 × 10−2 mm m−1 and 0.74 μm m−1, respectively. Mesophyll conductance (gm) and CO2 assimilation in ambient [CO2] (Aamb) decreased by 1.1 mmol m−2 s−1 per m and 0.082 μmol m−2 s−1 per m increase in height, respectively. Mean reductions in gm and Aamb of foliage from 5 to 55 m were 47% and 42%, respectively. The observed trend in Aamb was associated with gm and several leaf anatomic characteristics that are likely to be determined by the prevailing vertical tension gradient during foliar development. A linear increase in foliar δ13C values with height (0.042‰ m−1) implied that relative stomatal and mesophyll limitations of photosynthesis in intact shoots increased with height. These data suggest that increasing height leads to both fixed structural constraints on leaf gas exchange and dynamic constraints related to prevailing stomatal behavior.Keywords: leaf anatomy, mesophyll resistance, photosynthesis, growth limitationKeywords: leaf anatomy, mesophyll resistance, photosynthesis, growth limitatio

An Annual Pattern of Native Embolism in Upper Branches of Four Tall Conifer Species

Premise of the study: The Pacific Northwest of North America experiences relatively mild winters and dry summers. For the tall coniferous trees that grow in this region, we predicted that loss in the hydraulic conductivity of uppermost branches would be avoided because of difficulty reversing accumulated emboli in xylem that is always under negative pressure. • Methods: To test this hypothesis, we measured native percent loss in hydraulic conductivity (PLC; the decrease of in situ hydraulic conductivity relative to the maximum) monthly throughout 2009 in branches at the tops (~50 m) of four species in an old growth forest in southern Washington. • Key results: Contrary to our prediction, freeze – thaw cycles resulted in considerable native PLC. Branches showed hydraulic recovery in the spring and after a moderate increase in native embolism that was observed after an unusually hot period in August. The September recovery occurred despite decreases in the leaf and stem water potentials compared to August values. • Conclusions: Recoveries in branches of these trees could not have occurred by raising the water potential enough to dissolve bubbles simply by transporting water from roots and must have occurred either through water absorption through needles and/ or refilling under negative pressure. Excluding the August value, native embolism values correlated strongly with air temperature of the preceding 10 d. For three species, we found that branches with lower wood density had higher specific conductivity, but not greater native PLC than branches with higher wood density, which calls into question whether there is any hydraulic benefit to higher wood density in small branches in those species.Keywords: Tsuga heterophylla, Abies grandis, wood density, hydraulic conductivity, Thuja plicata, Pseudotsuga menziesi

Safety factors for xylem failure by implosion and air-seeding within roots, trunks and branches of young and old conifer trees

The cohesion-tension theory of water transport states that hydrogen bonds hold water molecules together and that they are pulled through the xylem under tension. This tension could cause transport failure in at least two ways: collapse of the conduit walls (implosion), or rupture of the water column through air-seeding. The objective of this research was to elucidate the functional significance of variations in tracheid anatomical features, earlywood to latewood ratios and wood densities with position in young and old Douglas-fir and ponderosa pine trees in terms of their consequences for the safety factors for tracheid implosion and air-seeding. For both species, wood density increased linearly with percent latewood for root, trunk and branch samples. However, the relationships between anatomy and hydraulic function in trunks differed from those in roots and branches. In roots and branches increased hydraulic efficiency was achieved at the cost of increased vulnerability to air-seeding. Mature wood of trunks had earlywood with wide tracheids that optimized water transport and had a high percentage of latewood that optimized structural support. Juvenile wood had higher resistance to air-seeding and cell wall implosion. The two safety factors followed similar axial trends from roots to terminal branches and were similar for both species studied and between juvenile and mature wood.Keywords: tracheid, cell wall, mature wood, juvenile wood, embolism, water transpor

The dynamic pipeline: hydraulic capacitance and xylem hydraulic safety in four tall conifer species

Recent work has suggested that plants differ in their relative reliance on structural avoidance of embolism versus maintenance of the xylem water column through dynamic traits such as capacitance, but we still know little about how and why species differ along this continuum. It is even less clear how or if different parts of a plant vary along this spectrum. Here we examined how traits such as hydraulic conductivity or conductance, xylem vulnerability curves, and capacitance differ in trunks, large- and small-diameter branches, and foliated shoots of four species of co-occurring conifers. We found striking similarities among species in most traits, but large differences among plant parts. Vulnerability to embolism was high in shoots, low in small- and large-diameter branches, and high again in the trunks. Safety margins, defined as the pressure causing 50% loss of hydraulic conductivity or conductance minus the midday water potential, were large in small-diameter branches, small in trunks and negative in shoots. Sapwood capacitance increased with stem diameter, and was correlated with stem vulnerability, wood density and latewood proportion. Capacitive release of water is a dynamic aspect of plant hydraulics that is integral to maintenance of long-distance water transport.This is the publisher’s final pdf. The published article is copyrighted by John Wiley & Sons Ltd. and can be found at: http://onlinelibrary.wiley.com/journal/10.1111/%28ISSN%291365-3040Keywords: vulnerability curves, safety margins, hydraulic conductance, conifers, capacitanc

Evidence for xylem embolism as a primary factor in dehydration-induced declines in leaf hydraulic conductance

Hydraulic conductance of leaves (K[subscript leaf]) typically decreases with increasing water stress and recent studies have proposed different mechanisms responsible for decreasing K[subscript leaf]. We measured K[subscript leaf] concurrently with ultrasonic acoustic emissions (UAEs) in dehydrating leaves of several species to determine whether declining K[subscript leaf] was associated with xylem embolism. In addition, we performed experiments in which the surface tension of water in the leaf xylem was reduced by using a surfactant solution. Finally, we compared the hydraulic vulnerability of entire leaves with the leaf lamina in three species. Leaf hydraulic vulnerability based on rehydration kinetics and UAE was very similar, except in Quercus garryana. However, water potentials corresponding to the initial decline in K[subscript leaf] and the onset of UAE in Q. garryana were similar. In all species tested, reducing the surface tension of water caused K[subscript leaf] to decline at less negative water potentials compared with leaves supplied with water. Microscopy revealed that as the fraction of embolized xylem increased, K[subscript leaf] declined sharply in Q. garryana. Measurements on leaf discs revealed that reductions in lamina hydraulic conductance with dehydration were not as great as those observed in intact leaves, suggesting that embolism was the primary mechanism for reductions in K[subscript leaf] during dehydration