Water relations of stem succulent trees in north-central Baja California

Summary Water relations of several stem succulent trees were measured in north-central Baja California in comparisons to other growth forms in the same habitat. Our research concentrated on three stem succulent species (Idria collumnaris, Pachycormus discolor and Bursera microphylla) each with a different succulent stem morphology. The stem succulent trees had 1 to 4 kg H₂O/m³ of trunk while the other trees and shrubs in the same habitat had 0.6 to 0.8 kg H₂O/m³. The diurnal and seasonal variation in leaf water potential was small for the stem succulent species in comparison to deciduous and evergreen species as a consequence of the stem-water, buffering capacity. In addition, the leaf conductance of the stem succulent species was low (60 mmol m^–2 s^–1) and yet, the leaf conductance decreased through the day similar to adjacent evergreen and deciduous species. The leaves of the stem succulent trees lost turgor at low saturated water deficits (0.06 to 0.14), had comparatively high osmotic potentials, and high values of elastic modulus in comparison to adjacent evergreen and deciduous species. The stem acts as an important buffering mechanism allowing for the maintenance of leaf turgor in these stem succulent trees. The low transpiration rates of the stem succulent trees may be a mechanism to minimize leaf saturated water deficit and extend leaf longevity.     

Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ_L) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although Ψ_L did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ¹⁸O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar ¹⁸O (∆¹⁸O_l) and ¹³C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.     

Does irrigation affect leaf phenology in deciduous and evergreen trees of the savannas of northern Australia?

Abstract Soil moisture was augmented experimentally during two successive dry seasons and the intervening wet season in a humid tropical savanna in Darwin, northern Australia. Leaf phenology was monitored in four common tree species Termmalia ferdinandiana and Planchonia careya (both deciduous species), and Eucalyptus miniata and Eucalyptus tetrodonta (both evergreen species). Irrigation produced consistently significant effects in only T. ferdinandiana. In this species leaf-flush was significantly earlier, canopy decline and leaf-fall were significantly later and the attainment of full canopy was earlier in irrigated compared with non-irrigated trees. Litterfall, and the seasonal patterns of contraction and expansion of stems (a measure of stem water status or storage) were not significantly affected by irrigation in any species. Leaf longevity in the deciduous species was 6–8 months; some eucalypt leaves lived for approximately 1 year, but none lived longer than 18 months. Irrigation had relatively little effect on leaf longevity. While variation in soil moisture is a potentially important cue to both leaf-fall and leaf-flush, stem water status and climatic factors such as vapour pressure deficit may also be important climatic cues to phenological behaviour.     

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

Phenology of temperate trees in tropical climates

Several North American broad-leaved tree species range from the northern United States at 47°N to moist tropical montane forests in Mexico and Central America at 15–20°N. Along this gradient the average minimum temperatures of the coldest month (T _Jan), which characterize annual variation in temperature, increase from –10 to 12°C and tree phenology changes from deciduous to leaf-exchanging or evergreen in the southern range with a year-long growing season. Between 30 and 45°N, the time of bud break is highly correlated with T _Jan and bud break can be reliably predicted for the week in which mean minimum temperature rises to 7°C. Temperature-dependent deciduous phenology—and hence the validity of temperature-driven phenology models—terminates in southern North America near 30°N, where T _Jan>7°C enables growth of tropical trees and cultivation of frost-sensitive citrus fruits. In tropical climates most temperate broad-leaved species exchange old for new leaves within a few weeks in January-February, i.e., their phenology becomes similar to that of tropical leaf-exchanging species. Leaf buds of the southern ecotypes of these temperate species are therefore not winter-dormant and have no chilling requirement. As in many tropical trees, bud break of Celtis, Quercus and Fagus growing in warm climates is induced in early spring by increasing daylength. In tropical climates vegetative phenology is determined mainly by leaf longevity, seasonal variation in water stress and day length. As water stress during the dry season varies widely with soil water storage, climate-driven models cannot predict tree phenology in the tropics and tropical tree phenology does not constitute a useful indicator of global warming.     

Coordinated changes in photosynthesis,water relations and leaf nutritional traits of canopy trees along a precipitation gradient in lowland tropical forest

We investigated leaf physiological traits of dominant canopy trees in four lowland Panamanian forests with contrasting mean annual precipitation (1,800, 2,300, 3,100 and 3,500 mm). There was near complete turn-over of dominant canopy tree species among sites, resulting in greater dominance of evergreen species with long-lived leaves as precipitation increased. Mean structural and physiological traits changed along this gradient as predicted by cost–benefit theories of leaf life span. Nitrogen content per unit mass (N_mass) and light- and CO₂-saturated photosynthetic rates per unit mass (P_mass) of upper canopy leaves decreased with annual precipitation, and these changes were partially explained by increasing leaf thickness and decreasing specific leaf area (SLA). Comparison of 1,800 mm and 3,100 mm sites, where canopy access was available through the use of construction cranes, revealed an association among extended leaf longevity, greater structural defense, higher midday leaf water potential, and lower P_mass, N_mass, and SLA at wetter sites. Shorter leaf life spans and more enriched foliar ¹⁵N values in drier sites suggest greater resorption and re-metabolism of leaf N in drier forest. Greater dominance of short-lived leaves with relatively high P_mass in drier sites reflects a strategy to maximize photosynthesis when water is available and to minimize water loss and respiration costs during rainless periods. Overall, our study links coordinated change in leaf functional traits that affect productivity and nutrient cycling to seasonality in lowland tropical forests.     

Plant diversity patterns in semi-natural grasslands along an elevational gradient in southern Norway

Leaf demography and growth of six common, co-occurring woody plant species that varied in stature (tree vs. shrub) and leaf texture (sclerophyllous, coriaceous, malacophyllous) were examined in a subtropical savanna parkland in southern Texas, USA. We tested the hypotheses that, (a) leaves of plants with evergreen canopies would have longer life spans than those of deciduous species; (b) supplementation of soil moisture would decrease leaf life span in both evergreen and deciduous species; (c) species responses to increased soil moisture availability would be inversely related to leaf longevity; and (d) deciduous growth forms would exhibit a greater growth response to increased soil moisture availability than their evergreen counterparts.A variety of seasonal leaf habits (evergreen, winter-deciduous and summer-deciduous canopies) and leaf life spans (median = 66 to 283 days) were represented by the targeted species, but there was no clear relationship between seasonal leaf habit and leaf longevity. Among species with evergreen canopies, median leaf longevity ranged from short (Zanthoxylum fagara = 116 days; Condalia hookeri = 158 days) to long (Berberis trifoliolata = 283 days) but did not exceed 1 yr. In fact, leaf longevity in evergreen shrubs was often comparable to, or shorter than, that of species with deciduous canopies (Ziziphus obtusifolia = 66 days; Diospyros texana = 119 days; Prosopis glandulosa = 207 days). Augmentation of surface soil moisture had no detectable effect on median leaf life span in any species and there was no clear relationship between leaf longevity and species growth responses to irrigation. Contrary to expectations, species with evergreen canopies responded to irrigation by producing more leaf biomass, longer shoots and more leaf cohorts/year than did deciduous species.Species differences in the annual cycle of leaf initiation, leaf longevity and canopy development, combined with contrasts in root distributions and a highly variable climate, may allow for spatial and temporal partitioning of resources and hence, woody species coexistence and diversity in this system. However, the lack of expected relationships between leaf longevity, leaf habit and plant responses to resource enhancement suggests that structure-function relationships and functional groupings developed in strongly seasonal environments cannot be applied with confidence to these subtropical savannas and thorn woodlands.     

Water chemistry and nutrient budgets in an undisturbed evergreen rainforest of southern Chile

In a pristine evergreen rainforest of Nothofagus betuloides, located at the Cordillera de los Andes in southern Chile (41 °S), concentrations and fluxes of nutrients in bulk precipitation, cloud water, throughfall water, stemflow water, soil infiltration and percolation water and runoff water were measured. The main objectives of this study were to investigate canopy-soil-atmosphere interactions and to calculate input-output budgets. From May 1999 till April 2000, the experimental watershed received 8121 mm water (86% incident precipitation, 14% cloud water), of which the canopy intercepted 16%. Runoff water volume amounted 9527 mm. Bulk deposition of inorganic (DIN) and organic (DON) nitrogen amounted 3.6 kg ha^–1 year^–1 and 8.2 kg ha^–1 year^–1 respectively. Occult deposition (clouds + fog) contributes for 40% to the atmospheric nitrogen input (bulk + occult deposition) of the forest. An important part of the atmospheric ammonium deposition is retained within the canopy or converted to nitrate or organic nitrogen by epiphytic bacteria or lichens. Also the export of inorganic (0.9 kg ha^–1 year^–1) and organic (5.2 kg ha^–1 year^–1) nitrogen via runoff is lower than the input to the forest floor via throughfall and stemflow water (3.2 kg DIN ha–1 year^–1 and 5.6 kg DON ha^–1 year^–1). The low concentrations of NO ₃ ^– and NH ₄ ⁺ under the rooting depth suggest an effective biological immobilization by vegetation and soil microflora. Dry deposition and foliar leaching of base cations (K⁺, Ca²⁺, Mg²⁺) was estimated using a canopy budget model. Bulk deposition accounted for about 50% of the total atmospheric input. Calculated dry and occult deposition are both of equal value (about 25%). Foliar leaching of K⁺, Ca²⁺, and Mg²⁺ accounted for 45%, 38% and 6% of throughfall deposition respectively. On an annual basis, the experimental watershed was a net source for Na⁺, Ca²⁺ and Mg²⁺.     

Aspects of tissue water relations and seasonal changes of leaf water potential components of evergreen and deciduous species coexisting in tropical dry forests

Summary This study compared the tissue water relations and seasonal changes in leaf water potential components of an evergreen tree,Morisonia americana, and two evergreen shrubs,Capparis verrucosa andC. aristiquetae, with two deciduous trees,Humboltiella arborea andLonchocarpus dipteroneurus, and the deciduous vineMansoa verrucifera. All these species coexist in a tropical dry forest in Venezuela. Leaves of the evergreen species are sclerophyllous, while those of the deciduous species are mesophytic. Leaf area to leaf weight ratios of fully mature leaves were about 75 and 170 cm² g^–1 in evergreen and deciduous species, respectively. Seasonal fluctuations of leaf water content per unit of dry weight, water potential, and turgor pressure were smaller in evergreen than in deciduous species. The analysis of tissue water relations using pressurevolume curves showed that evergreen species could develop a higher leaf turgor and lose turgor at lower leaf water potentials than deciduous species. This was related to a lower osmotic potential at full turgor in evergreen (-3.0 MPa)_than in deciduous (-2.0 MPa) species, rather than to the elastic properties of leaf tissue. The volumetric modulus of elasticity was 14 MPa in evergreen compared with 7–10 MPa in deciduous species. Thus, leaf characteristics are important in determining the drought resistance of evergreen species of this tropical dry forest.     

Regional variation in canopy transpiration of Central European beech forests

Forest hydrologists have hypothesised that canopy transpiration (E_c) of European temperate forests occurs at rather similar rates in stands with different tree species and hydrologic regimes. We tested this hypothesis by synchronously measuring xylem sap flow in four mature stands of Fagus sylvatica along a precipitation gradient with the aim (1) of exploring the regional variability of annual canopy transpiration (E_c(t)) in this species, and (2) of analysing the relationship between precipitation (P) and E_c(t). E_c(t) rates of 216, 225, 272 and 303 mm year^–1 corresponded to precipitation averages of 520, 710, 801 and 1,040 mm year^–1 in the four stands. We explored the regional variability of E_c(t) in Central European colline to sub-montane beech stands in two meta-analyses based on (1) existing sap flow data on beech (n=5 observations), or (2) all canopy transpiration data on beech obtained by different techniques (sap flow, micrometeorological or soil water budget approaches, n=25). With a coefficient of variation (CV) of 20%, the regional variability of E_c(t) (213–421 mm year^–1) was smaller than the variation in corresponding precipitation (550–1,480 mm year^–1). The mean E_c(t) for beech was 289 (±58) mm year^–1 (n=25). A humped-shaped relationship between E_c(t) and P, with a broad transpiration maximum in the precipitation range from ca. 700 to 1,000 mm year^–1, was found which may indicate soil moisture limitation of transpiration for P –1, and reduced transpiration by increased cloudiness or leaf wetness for P>1,000 mm year^–1. Thus, the precipitation level significantly influences canopy transpiration of humid temperate forests; however, the size of the P influence on E_c(t) and, in part, the direction of its effect differ from forests in semi-arid or arid climates. European beech has the capacity to maintain high E_c rates in both humid and partly dry summer climates (P<550 mm year^–1).     

Induction of rehydration and bud break by irrigation or rain in decidous trees of a tropical dry forest in Costa Rica

Summary Clusters of 2–4 bare, deciduous hardwood trees and woody vines in a dry upland forest in Costa Rica were surrounded by scaffolding and rehydration was induced during the dry season by irrigation of 9–50 m² plots with 200 mm water. The resulting changes in water status preceding bud break were monitored. Following irrigation, stem water potentials increased from < –4 MPa to about –1.5 MPa within 24 h and to > –0.3 MPa within 48 h. Rehydration of stem tissues by lateral transport, measured as an increase in electric conductivity, continued for 4–8 days. Terminal flower buds in Tabebuia ochracea began to expand 48 h after irrigation and trees were in full bloom 4 days later. In all experimental species, lateral vegetative buds began to expand 5–7 days after irrigation and leaves were fully expanded 2 weeks later. After the first rains of the rainy season (100 mm in 48 hr) all trees in the dry forest rehydrated and leaves emerged in synchrony slightly faster than after irrigation. In response to rain or irrigation drought-stressed tropical hardwood trees thus rehydrated at rates similar to those of desert succulents and their development resumed much faster than that of deciduous cold-temperate trees in spring.     

Seasonal variations of leaf water potential and growth in fertigated carob-trees (Ceratonia siliqua L.)

Variations of predawn and midday leaf water potential and relative growth rates were studied in mature carob trees (Ceratonia siliqua L. cv Mulata) submitted to a fertigation experiment. Three levels of irrigation were tested: 0%, 50% and 100%, based on daily standard evaporation values. For each irrigation level two nitrogen amounts were applied –21 and 63 kg N ha^-1 year^-1 as ammonium nitrate. The experiment was run between July 91 and August 1993. Measurements of leaf water potential and absolute branch length increments were made at monthly intervals, during the entire experimental period or during seasonal growth, respectively. Leaf water potential was related to soil volumetric water content, maximum and minimum air temperature and daily evaporation. Predawn leaf water potentials were always higher than –1.1 MPa. Midday leaf water potential values presented very large seasonal variations and very low values independent of treatments. The low leaf water potentials observed for the fertigated trees during summer, suggest that this parameter may be related not only to the evaporative demand but also to growth investment. The amount of fertigation was positively correlated with vegetative growth increment and fruit production. Practical implications for irrigation schedules of leaf water potential patterns together with drought adaptation mechanisms of carob tree are discussed.     

Altitudinal distribution of evergreen broad-leaved trees and their leaf-size pattern on a humid subtropical mountain,Mt. Emei,Sichuan, China

Altitudinal distribution of evergreen broad-leaved trees and changes in their leaf sizes were studied on a humid subtropical mountain, Mt. Emei (3099 m a.s.l., 29°34.5 N, 103°21.5 E), Sichuan, China. Among the total woody flora of ca. 540 species, evergreen broad-leaved trees account for 88 species in 39 genera and 23 families, corresponding to the northern limit of subtropical evergreen broad-leaved trees. The number of evergreen broad-leaved tree species greatly decreased from the low-altitudinal, evergreen broad-leaved forest zone (600–1500 m) to the mid-altitudinal, mixed forest zone (1500–2500 m), and to the high-altitudinal, coniferous forest zone (2500–3099 m). The overall trend of reduced leaf size toward upper zones was analyzed and documented in detail. The 88 species were assigned to three leaf-size classes: notophylls (48%), microphylls (36%), and mesophylls (16%). The leaf size was relatively small and the specific leaf weight (SLW, mg cm^–2) was much larger in high altitude as compared to low altitude. No overall correlation was found between leaf size and SLW, but leaf size decreased as SLW increased toward high altitude for certain species having relatively wide altitudinal ranges. Moreover, leaf size varied with forest stratification: canopy trees were predominantly notophyllous species, while subcanopy and understorey trees were mainly microphyllous species. The tendency is compatible with the trend found in other mountains of East Asia.     

Field water relations of three temperate vines

Summary A two year comparative field water relations study was conducted in central Maryland on three sympatric temperate lianas, Lonicera japonica, Vitis vulpina and Parthenocissus quinquefolia. Seasonal physiological activity was longer by approximately 9 weeks in the evergreen L. japonica, while peak rates of stem elongation were 4–10 fold higher in the two deciduous species. There were marked differences in vascular anatomy and water use patterns among the three species, however all three evidenced varying degrees of stomatal closure in response to decreasing soil water availability and increasing atmospheric evaporative demand. The range of leaf water potentials measured in these species was quite narrow in comparison to other temperate woody species. Two of the species showed no alterations in their tissue water release properties in response to decreased soil water availability, while V. vulpina showed a limited capacity in this regard. Most significant among the species differences in water relations were the conservative water use patterns of P. quinquefolia, and the midday maxima of transpirational water loss measured in L. japonica compared to the morning peaks in traspiration for the two deciduous species. The differences found in anatomy, leaf phenology, climbing mechanics, water relations and canopy development among these three sympatric vines implies a spatial and temporal partitioning of light and water resources and emphasizes the diversity of morphological-physiological suites of characters present among species co-occurring in the same macrohabitat.     

Differential impact of liana colonization on the leaf functional traits of co-occurring deciduous and evergreen trees in a tropical dry scrub forest

The present study was carried out to analyze the leaf functional traits of co-occurring evergreen and deciduous tree species in a tropical dry scrub forest. This study also intended to check whether the species with contrasting leaf habits differ in their leaf trait plasticity, responding to the canopy infestation by lianas. A total of 11 leaf functional traits were studied for eight tree species with contrasting leaf habits (evergreen and deciduous) and liana-colonization status (with or without liana). In the liana-free environment (L^–), evergreen trees had significantly higher leaf tissue density (LTD) and total chlorophyll (CHL_t) than the deciduous species. Whereas the deciduous trees had higher specific leaf area (SLA) and mass-based leaf nitrogen concentration (N_mass). The leaf trait-pair relationship in the present study agreed with the well-established global trait-pair relationships (leaf thickness (LT) vs. SLA, N_mass vs. LT, SLA vs. N_mass, and LDMC vs. SLA). There was a significant difference between L⁺ and L^– individuals in leaf area (LA), petiole length (PL), SLA, LDMC, and CHL_t in the deciduous species. On the other hand, evergreen species showed marked differences across LT, SLA, LTD, N_mass, and chlorophyll components between L⁺ and L^– individuals of the same species. The results revealed the differential impact of liana colonization on the host trees with contrasting leaf habits. The deciduous species with the acquisitive strategy can have a competitive advantage over evergreen species in the exposed environments (L^–), whereas evergreen species with shade-tolerant properties were better acclimated to the shaded environments (L⁺). Therefore, liana colonization can significantly impact the C-fixation strategies of the host trees by altering their light environment and further, the magnitude of such impact may vary among species of different leaf habits. The result also indicated the patterns of convergence and divergence in some of the leaf functional traits between evergreen and deciduous species explaining the patterns of species co-existence.

     

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements

Summary Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day^–1 tree^–1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m² of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s^–1 that is equivalent to 112 mmol m^–2 s^–1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m^–2 s^–1 at the top and 150 mmol m^–2 s^–1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was –27.76±0.27 (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.This paper is dedicated to Prof. Dr. O.L. Lange on the occasion of his 65th birthday     

Environmental control of canopy dynamics and photosynthetic rate in the evergreen tussock grass Stipa tenacissima

Seasonal changes in leaf demography and gas exchange physiology in the tall evergreen tussock grass Stipa tenacissima, one of the few dominant plant species in the driest vegetation of Europe, were monitored over a period of two years at a field site in semi-arid south-eastern Spain. Three age-classes of leaves – young, mature and senescent – were distinguished in the green canopy. Production of new leaves and extension growth of older leaves occurred exclusively from October–November to May–June. The rate of extension was significantly correlated with gravimetric soil water content. Leaf growth ceased after gravimetric soil water content fell below 0.015 g g^–1 at the beginning of the dry season which corresponded to pre-dawn leaf water potentials of -3.0 MPa. Leaf senescence and desiccation reduced green leaf area by 43–49% during the dry season. Diurnal changes in the net photosynthetic rate of all three cohorts of leaves were bimodal with an early morning maximum, a pronounced midday depression and a small recovery late in the afternoon. Maximum photosynthetic rates of 10–16 mol CO₂ m^–2 s^–1 were attained from November 1993 to early May 1994 in young and mature leaves. Photosynthetic rate declined strongly during the dry season and was at or below compensation in September 1994. Gas exchange variables of young and mature leaves were not significantly different, but photosynthetic rate and diffusive conductance to water vapour of senescing leaves were significantly lower than in the two younger cohorts. Leaf nitrogen content of mature leaves varied seasonally between 2.9 and 5.2 g m^–2 (based on projected area of folded leaves), but was poorly correlated with maxima of the photosynthetic rate. There was a stronger linear relationship between the daily maxima of leaf conductance and pre-dawn leaf water potential than with atmospheric water vapour saturation deficit. Seasonal and between-year variation in daily carbon assimilation were caused mainly by differences in climatic conditions and canopy size whereas the effect of age structure of canopies was negligible. Since water is the most important limiting factor for growth and reproduction of S. tenacissima, any future rise in mean temperature, which might increase evapotranspiration, or decrease in rainfall, may considerably reduce the productivity of the grasslands, particularly at the drier end of their geographical distribution.     

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2–4-mm diameter) collected at 20–30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance (g_s) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20–30 cm depth above –0.5 MPa in the old-growth Douglas-fir site and –1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about –2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g_s decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.     

Leafing patterns and leaf traits of four evergreen shrubs in the Patagonian Monte, Argentina

We assessed leafing patterns (rate, timing, and duration of leafing) and leaf traits (leaf longevity, leaf mass per area and leaf-chemistry) in four co-occurring evergreen shrubs of the genus Larrea and Chuquiraga (each having two species) in the arid Patagonian Monte of Argentina. We asked whether species with leaves well-defended against water shortage (high LMA, leaf longevity, and lignin concentration, and low N concentration) have lower leaf production, duration of the leafing period, and inter-annual variation of leafing than species with the opposite traits. We observed two distinctive leafing patterns each related to one genus. Chuquiraga species produced new leaves concentrated in a massive short leafing event (5–48 days) while new leaves of Larrea species emerged gradually (128–258 days). Observed leafing patterns were consistent with simultaneous and successive leafing types previously described for woody plants. The peak of leaf production occurred earlier in Chuquiraga species (mid September) than in Larrea species (mid October–late November). Moreover, Chuquiraga species displayed leaves with the longest leaf lifespan, while leaves of Larrea species had the lowest LMA and the highest N and soluble phenolics concentrations. We also observed that only the leaf production of Larrea species increased in humid years. We concluded that co-occurring evergreen species in the Patagonian Monte displayed different leafing patterns, which were associated with some relevant leaf traits acting as plant defenses against water stress and herbivores. Differences in leafing patterns could provide evidence of ecological differentiation among coexisting species of the same life form.     

Dynamics of leaf minerals,bdleaf area,and biomass from hardwoods intensively grown on a peat bog

Summary Water willow and grey alder were grown on a raised sphagnum bog in central Sweden. The stands were intensively treated by daily irrigation and fertilization during the growing period in order to improve site fertility. After a 2-year establishment period high production rates were achieved in willow stands, 0.8 kg stem dry weight m^–2 year^–1 on current plus one (C+1) year old shoots. In these stands the canopy was closed with a leaf area index (LAI) that peaked at approximately 7. The canopy in the alder stand did not close during the initial 3 years of growth and the measured production rate was relatively low, at approximately 0.4 kg dry weight m^–2 year ^–1 in the last year. The leaf nitrogen content was 3%–4% of dry weight during the summer and the other studied mineral elements were in almost optimal proportion to nitrogen. This was considered to be an effect of the intensive fertilization regime. Above-ground production close to maximum yield was attained in the prevailing conditions of soil, climate and biomass partitioning.