Effect of moisture availability on photosynthetic productivity and autotrophic respiration of an oak stand

The gas exchange of an oak stand in the southeastern forest steppe was examined under variable moisture availability and ambient temperature. For several years the CO₂ exchange was measured concurrently on leafy shoots and nonphotosynthesizing organs (trunks, branches, and roots) of oak trees (Quercus robur L.). The root respiration was found to decrease upon moisture limitation. The trunk respiration was promoted by the increase in moisture deficiency until the predawn leaf water potential (PLWP) reduced to −1.5 MPa, and respiration declined gradually upon further reduction in moisture availability. The proportion between net and gross primary productivities (NPP/GPP) also depended on moisture availability. The NPP/GPP ratio was 0.49 under optimal conditions at PLWP of −0.5 MPa, but it approached zero when PLWP lowered to −1.5 MPa. Upon drought conditions, which are typical of southeastern steppe in the second half of summer, the oak stand lost its CO₂ sink capacity and became the CO₂ source.     

In situ measurement of fine root water absorption in three temperate tree species—Temporal variability and control by soil and atmospheric factors

Miniature heat balance-sap flow gauges were used to measure water flows in small-diameter roots (3–4 mm) in the undisturbed soil of a mature beech–oak–spruce mixed stand. By relating sap flow to the surface area of all branch fine roots distal to the gauge, we were able to calculate real time water uptake rates per root surface area (J_s) for individual fine root systems of 0.5–1.0 m in length. Study aims were (i) to quantify root water uptake of mature trees under field conditions with respect to average rates, and diurnal and seasonal changes of J_s, and (ii) to investigate the relationship between uptake and soil moisture θ, atmospheric saturation deficit D, and radiation I. On most days, water uptake followed the diurnal course of D with a mid-day peak and low night flow. Neighbouring roots of the same species differed up to 10-fold in their daily totals of J_s (<100–2000 g m⁻² d⁻¹) indicating a large spatial heterogeneity in uptake. Beech, oak and spruce roots revealed different seasonal patterns of water uptake although they were extracting water from the same soil volume. Multiple regression analyses on the influence of D, I and θ on root water uptake showed that D was the single most influential environmental factor in beech and oak (variable selection in 77% and 79% of the investigated roots), whereas D was less important in spruce roots (50% variable selection). A comparison of root water uptake with synchronous leaf transpiration (porometer data) indicated that average water fluxes per surface area in the beech and oak trees were about 2.5 and 5.5 times smaller on the uptake side (roots) than on the loss side (leaves) given that all branch roots <2 mm were equally participating in uptake. Beech fine roots showed maximal uptake rates on mid-summer days in the range of 48–205 g m⁻² h⁻¹ (i.e. 0.7–3.2 mmol m⁻² s⁻¹), oak of 12–160 g m⁻² h⁻¹ (0.2–2.5 mmol m⁻² s⁻¹). Maximal transpiration rates ranged from 3 to 5 and from 5 to 6 mmol m⁻² s⁻¹ for sun canopy leaves of beech and oak, respectively. We conclude that instantaneous rates of root water uptake in beech, oak and spruce trees are above all controlled by atmospheric factors. The effects of different root conductivities, soil moisture, and soil hydraulic properties become increasingly important if time spans longer than a week are considered.     

Leaf productivity along a precipitation gradient in lowland Panama: patterns from leaf to ecosystem

Moisture availability has the potential to affect tropical forest productivity at scales ranging from leaf to ecosystem. We compared data for leaf photosynthetic, chemical and structural traits of canopy trees, litterfall production and seasonal availability of soil water at four sites across a precipitation gradient (1,800–3,500 mm year^–1) in lowland Panamanian forest to determine how productivity at leaf and ecosystem scales may be related. We found stronger seasonality in soil water potential at drier sites. Values were close to zero at all sites during the wet season and varied between a minimum of –2.5 MPa and –0.3 MPa at the driest and wettest sites, respectively, during the dry season. Leaf photosynthesis and nitrogen concentration decreased with increasing precipitation, whereas leaf thickness increased with increasing precipitation. Leaf toughness and fiber/N ratios increased with increasing precipitation indicating reduced nutritional content and palatability with precipitation. Seasonality of litter production and quality decreased with increasing precipitation, but the amount of litterfall produced was not substantially different among sites. It appears that in Neotropical forest, moisture availability is associated with leaf photosynthetic and defensive traits that influence litterfall timing and quality. Therefore, variation in leaf physiological traits has the potential to influence decomposition and nutrient cycling through effects on litter quality.     

State and Phase Transition Behaviors ofQuercus rubraSeed Axes and Cotyledonary Tissues: Relevance to the Desiccation Sensitivity and Cryopreservation of Recalcitrant Seeds

Freezing and melting transitions of cellular water in embryonic axes and cotyledonary tissues of recalcitrantQuercus rubra(red oak) seeds were compared under slow and rapid cooling conditions. The relevance of desiccation sensitivity (critical water content) and state/phase transition behaviors to cryopreservation was examined. Under a slow to intermediate cooling condition (≤10°C min⁻¹), unfrozen water content in the tissues decreased to less than the critical water content, resulting in a dehydration damage. Under a rapid cooling condition (>100°C min⁻¹) using liquid nitrogen (LN₂), freeze-induced dehydration damage could be avoided if the initial water content was >0.50 g g⁻¹dry wt. However, at water content >0.50 g g⁻¹dry wt, the vitrified cellular matrix was highly unstable upon warming at 10°C min⁻¹. These results offered a theoretical explanation on the difficulty for successful cryopreservation of recalcitrant red oak embryonic axes. A complete state/phase transition diagram for red oak axes was constructed, and a vitrification-based cryopreservation protocol that employed predehydration and rapid cooling was examined. State/phase transition behaviors of cellular water are important parameters for cryopreservation; however, vitrification alone was not sufficient for seed tissues to survive the cryopreservation condition.     

灌溉量对苹果园内紫花苜蓿生物量与生理特征的影响

以苹果园内2年生紫花苜蓿为材料,设0.150、0.225(正常水平)和0.300m3/m2共3个灌溉量处理,研究不同灌溉量对紫花苜蓿生物量、光合生理指标、叶绿素含量、脯氨酸含量及电导率的影响。结果显示:(1)灌溉量对紫花苜蓿地上生物量有显著的影响,且随灌溉量的增加而增加;灌溉量大的处理苜蓿吸收根分布有浅层化趋势。(2)不同灌溉量处理的紫花苜蓿叶片净光合速率(Pn)与光合有效辐射(PAR)的变化趋势基本一致,均表现为在低PAR时Pn迅速增加,达到一定PAR后,Pn增幅渐趋平缓;净光合速率均随灌水量的增加而增加。(3)灌溉后土壤含水率和苜蓿叶片叶绿素含量随灌溉量的增加而升高,苜蓿叶片相对电导率随着灌水量的增加而呈现降低的趋势,低灌溉量可导致紫花苜蓿叶片中脯氨酸含量大量积累。研究表明,适当的灌溉可以有效改善苹果园土壤水分状况,提高紫花苜蓿光合生产力。     

Tree structure,regeneration and woody biomass removal in a sub-tropical forest of Mamlay watershed in the Sikkim Himalaya

This paper reports on the tree structure, tree dimension relationships and woody biomass production and removal of a sub-tropical natural forest in the Mamlay watershed of the Sikkim Himalaya. The forest provides fuel, fodder and timber to four villages. Only 11 tree species were found growing in the tree stratum despite the high diversity in the stand (32 tree species). The forest shows good regeneration potential with 5474 seedlings/ha and 1776 saplings/ha, but the population structure revealed a marked paucity of trees of higher diameter classes due to removal of trees of lower diameters. Standing wood biomass of 362 Mg/ha is mainly shared by 4 dominating species in the stand. The boles are removed mainly for timber and fuel purposes and about 22 Mg/ha wood biomass was removed in between 1987–1991. Net Primary productivity of woody biomass of the forest is recorded to be 18 Mg/ha/year. 3.85 Mg/ha of annual woody biomass production was removed in the form of tree boles apart from lopping of branches.     

Bottlenecks to water transport in Quercus robur L.: the abscission zone and its physiological consequences

Cladoptosis, the abscission of twigs, is the main mechanism of changes in crown structure in senescing pedunculate oak (Quercus robur L.). We tested the hypotheses that abscission zones in nodes of old pedunculate oak trees reduce leaf-specific hydraulic conductance of shoots and thereby limit the stomatal conductance and assimilation.Hydraulic conductance and leaf-specific hydraulic conductance, measured with a high pressure flowmeter in 0.5–1.5 m long shoots, were significantly lower in shoots of low vigour compared to vigorous growing shoots in a 165-years-old stand in the southeast of Germany. Two types of bottlenecks to water transport could be identified in shoots of old oak trees, namely nodes and abscission zones. In young twigs, vessel diameter and vessel density in nodes with abscission zones were significantly reduced compared with internodes. In nodes without abscission zones, vessel density was significantly reduced. The reduction of hydraulic conductance was especially severe in the smallest and youngest shoots with diameters less than 2 mm. Internodes of 1–5 mm sapwood diameter had an average hydraulic conductance of 7.13×10⁻⁶±0.2×10⁻⁶ kg s⁻¹ m⁻¹ MPa⁻¹, compared to 4.54×10⁻⁶±0.3×10⁻⁶ kg s⁻¹ m⁻¹ MPa⁻¹ in those with nodes.Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf-specific hydraulic conductance. Maximum rate of net photosynthesis A_max of the most vigorous shoots (VC0) (7.34±0.55 μmol m⁻² s⁻¹) was significantly higher (P<0.001) than in shoots of other vigour classes (5.97±0.28 μmol m⁻² s⁻¹). Our data support the hypothesis that the changes in shoot and consequently crown architecture that are observed in ageing and declining trees can limit photosynthesis by reducing shoot hydraulic conductance. Abscission zones increase the hydraulic disadvantage of less vigorous compared to vigorously growing twigs. Cladoptosis might serve as a mechanism of selection between twigs of different efficiency.     

Shedding of older needle age classes does not necessarily reduce photosynthetic primary production of Norway spruce

The hypothesis that the frequently observed shedding of older needle age classes in stands of Norway spruce suffering from atmospheric pollutants has minimal effect on photosynthetic primary production was assessed. Using structural parameters of young Norway spruce [Picea abies (L.) Karst.] trees, an existing 3-dimensional canopy photosynthesis model was adapted and validated for conifer canopies. This model was employed to quantitatively demonstrate that, depending on the stand density, the loss in photosynthetic active foliage area may be compensated by higher photosynthetic rates of the remaining younger foliage age classes due to higher levels of photosynthetically active radiation (PAR) within the thinned tree crowns. Under certain circumstances, beneficial effects on the tree's water use efficiency may be expected. Extrapolating the model results to mature tree stands is addressed. The potential effects of higher PAR levels on abundance and species diversity of the epiphytic and understory vegetation, as well as the general importance of the older foliage age classes for spruce trees on a long term scale, are discussed.     

Spatial–temporal variation in soil respiration in an oak–grass savanna ecosystem in California and its partitioning into autotrophic and heterotrophic components

The spatial upscaling of soil respiration from field measurements to ecosystem levels will be biased without studying its spatial variation. We took advantage of the unique spatial gradients of an oak–grass savanna ecosystem in California, with widely spaced oak trees overlying a grass layer, to study the spatial variation in soil respiration and to use these natural gradients to partition soil respiration according to its autotrophic and heterotrophic components. We measured soil respiration along a 42.5 m transect between two oak trees in 2001 and 2002, and found that soil respiration under tree canopies decreased with distance from its base. In the open area, tree roots have no influence on soil respiration. Seasonally, soil respiration increased in spring until late April, and decreased in summer following the decrease in soil moisture content, despite the further increase in soil temperature. Soil respiration significantly increased following the rain events in autumn. During the grass growing season between November and mid-May, the average of CO₂ efflux under trees was 2.29 μmol m⁻² s⁻¹, while CO₂ efflux from the open area was 1.40 μmol m⁻² s⁻¹. We deduced that oak root respiration averaged as 0.89 μmol m⁻² s⁻¹, accounting for 39% of total soil respiration (oak root + grass root + microbes). During the dry season between mid-May and October, the average of CO₂ efflux under trees was 0.87 μmol m⁻² s⁻¹, while CO₂ efflux from the open areas was 0.51 μmol m⁻² s⁻¹. Oak root respiration was 0.36 μmol m⁻² s⁻¹, accounting for 41% of total soil respiration (oak root + microbes). The seasonal pattern of soil CO₂ efflux under trees and in open areas was simulated by a bi-variable model driven by soil temperature and moisture. The diurnal pattern was influenced by tree physiology as well. Based on the spatial gradient of soil respiration, spatial analysis of crown closure and the simulation model, we spatially and temporally upscaled chamber measurements to the ecosystem scale. We estimated that the cumulative soil respiration in 2002 was 394 gC m⁻² year⁻¹ in the open area and 616 gC m⁻² year⁻¹ under trees with a site-average of 488 gC m⁻² year⁻¹.     

马占相思林冠层气孔导度对环境驱动因子的响应

利用Granier热消散探针在2003年10月测定了广东鹤山丘陵地马占相思林14株样树的树干液流,同时监测林冠上方的光合有效辐射、空气湿度和气温,结合树木的形态和林分的结构特征,计算马占相思的整树蒸腾(E)、林分总蒸腾(E_t)以及冠层平均气孔导度(g_c),分析树形特征与整树水分利用的关系、冠层气孔导度对光合有效辐射(PAR)和空气水汽压亏缺(D)的响应.结果表明,整树蒸腾与胸径(P＜0.0001)、边材面积(P＜0.0001)和冠幅(P=0.0007)以自然对数的形式、与树高(P=0.014)以幂函数的形式呈现显著正相关.冠层气孔导度最大值(g_cmax)随D的上升呈对数函数下降(P＜0.0001),对光合有效辐射的响应则呈双曲线函数增加(P＜0.0001).液流测定系统能提供连续和准确的整树和林分蒸腾速率值,经严格数学推导公式计算,最终可求出冠层气孔导度,是研究森林水分利用与环境因子相互关系的有效方法.     

Aquatic metabolism as an indicator of the ecological effects of hydrologic pulsing in flow-through wetlands

Water column metabolism is a major component in the functioning of wetland ecosystems and can be used as an indicator of ecosystem health. The effect of hydrologic pulsing on water column metabolism was studied with 2 year's field data and a validated model for two 1-ha created riparian wetlands in Columbus, OH, USA. Aquatic gross primary productivity (GPP) was measured during hydrologic pulses the first week of April, May and June of 2004 and compared to GPP during steady flow-conditions in April, May and June 2005. Pulses reduced diurnal variation of water temperature, pH and dissolved oxygen, and negatively affected GPP rates. Mean GPP measured during hydrologic flood pulses was 5.4 ± 2.6 kcal m⁻² day⁻¹, significantly lower than that measured for comparable months with steady-flow hydrology (10.8 ± 3.3 kcal m⁻² day⁻¹). Solar-normalized productivity values of 0.08 ± 0.01% of solar energy during pulses and 0.2 ± 0.02% for steady-flow conditions were also significantly different. Different hyperbolic curves of optimum productivity with water temperature were seen for pulsing and steady-flow conditions. A simulation model with hydrology, metabolism, and dissolved oxygen sub models was calibrated with 2005 steady-flow year data and validated with 2004 pulse year data. Results from both the field study and model simulations suggest that there was a threshold hydraulic inflow rate between 30 and 50 cm day⁻¹ where aquatic metabolism became negatively affected by flow.     

Effects of soil management practices and irrigation on plant water relations and productivity of chestnut stands under Mediterranean conditions

The effects of different soil management practices and irrigation on plant water relations, physiological response and productivity of chestnut stands in Northeastern Portugal were assessed during four growing seasons (2003 to 2006). Treatments were: conventional soil tillage up to 15–20 cm depth with a tine cultivator thrice a year (CT); no tillage with spontaneous herbaceous vegetation (NV); no tillage with rainfed seeded pasture (NP); and no tillage with irrigated seeded pasture (NIP). Results suggest that soil water availability was the most critical parameter for chestnut productivity over the study period. In all treatments, high predawn leaf water potentials (-0.40 to -0.55 MPa) were observed during the dry seasons of 2003, 2004 and 2006, showing no critical conditions for plant productivity, which is ascribed to water availability in deep soil layers. In contrast, in 2005, an extremely dry year, water potentials decreased and varied from -1.46 to -1.72 MPa in late summer, showing unfavourable conditions for nut production. Maintenance of spontaneous herbaceous vegetation without irrigation enhanced productivity of chestnut stands as compared with the conventional tillage system and the no tillage system with seeded pasture. Productivity in the soil watering system (NIP treatment) was not significantly different from that observed in the NV treatment. Therefore, studies on the irrigation strategy should be developed, in order to increase its efficiency especially in stands with young trees.     

Soil nitrogen dynamics in a holm oak forest

Soil nitrogen (N) dynamics were studied in a dense, holm oak (Quercus ilex ssp. ilex) stand in the Montseny mountains to determine annual and seasonal patterns of N availability and uptake in an undisturbed Mediterranean forest on acidic soil. Soil mineral N content, net N mineralization (NNM), and net nitrification (NN) were determined by monthly sampling at two soil depths followed by in situ incubation in polyethylene bags. NNM per unit of soil mass was much higher at 0–5 cm than at 5–20 cm (annual means 24 and 2.5 mg N/kg, respectively) but on an area basis NNM was similar at both depths. A total of 80 kg N/ha/yr were mineralized from the first 20 cm of soil. NN amounted to only 9% of the annual NNM (7.5 kg N/ha/yr) and it occurred only in the upper 5 cm. NNM was maximum in June and July, while the NN peaked in May. Despite favourable soil temperature and moisture, NNM was negative in autumn because of microbial immobilization. Seasonal and depth variations of NNM appeared to be controlled more by substrate quality than by organic matter quantity, temperature or moisture. NN was not limited by ammonium availability. Calculated N uptake amounted to 91 kg/ha yr, peaking in June and July. The investigated stand showed a moderately high N availability, but ammonium was the major form of mineral N supply for holm oak.     

Effects of forest successional status on microenvironmental conditions,diversity, and distribution of filmy fern species in a temperate rainforest

The vertical distribution of Hymenophyllaceae species has been related to microenvironmental variations around host trees. We addressed the questions: Do the vertical microenvironmental conditions within forest stands of differing successional statuses vary significantly? Does the diversity of Hymenophyllaceae species differ between forest successional statuses? Are the vertical distribution and diversity of Hymenophyllaceae species related more to humidity or light availability? Are there any interspecific differences in the desiccation tolerance of these species which can be related to their vertical distribution? We characterized the microhabitat conditions (vapor pressure deficit [VPD], air relative humidity [RH], and light availability [PAR]) and the vertical distribution of Hymenophyllaceae species in host trees, in both a secondary forest and an old‐growth temperate rainforest in Chile. Chlorophyll fluorescence was used to monitor the integrity of the photosynthetic apparatus during desiccation experiments. The stand basal area, tree height, and leaf area index were all significantly greater in the old‐growth forest stands, but VPD, RH, and PAR showed no significant differences between the two forests. Both successional statuses showed the same amount of filmy fern species in terms of both abundance and diversity. In both successional statuses VPD and RH decreased while PAR increased with the height of the hosts. Regardless of the forest's age, abundance and diversity of filmy ferns were greater in microsites of greater humidity and less light availability. Desiccation tolerance differed significantly among Hymenophyllaceae species. The distribution pattern could be better explained by the specific microenvironmental requirements and desiccation tolerance rather than the forest's successional status.     

A model of stomatal conductance to quantify the relationship between leaf transpiration, microclimate and soil water stress

A model of stomatal conductance was developed to relate plant transpiration rate to photosynthetic active radiation (PAR), vapour pressure deficit and soil water potential. Parameters of the model include sensitivity of osmotic potential of guard cells to photosynthetic active radiation, elastic modulus of guard cell structure, soil‐to‐leaf conductance and osmotic potential of guard cells at zero PAR. The model was applied to field observations on three functional types that include 11 species in subtropical southern China. Non‐linear statistical regression was used to obtain parameters of the model. The result indicated that the model was capable of predicting stomatal conductance of all the 11 species and three functional types under wide ranges of environmental conditions. Major conclusions included that coniferous trees and shrubs were more tolerant for and resistant to soil water stress than broad‐leaf trees due to their lower osmotic potential, lignified guard cell walls, and sunken and suspended guard cell structure under subsidiary epidermal cells. Mid‐day depression in transpiration and photosynthesis of pines may be explained by decreased stomatal conductance under a large vapour pressure deficit. Stomatal conductance of pine trees was more strongly affected by vapour pressure deficit than that of other species because of their small soil‐to‐leaf conductance, which is explainable in terms of xylem tracheids in conifer trees. Tracheids transport water by means of small pit‐pairs in their side walls, and are much less efficient than the end‐perforated vessel members in broad‐leaf xylem systems. These conclusions remain hypothetical until direct measurements of these parameters are available.     

Vegetative and reproductive responses,oil yield and composition from olive trees (Olea europaea) under contrasting water availability during the dry winter‐spring period in central Argentina

In Argentina, the climatic pattern of the olive production areas is characterised by a marked water deficit during winter and spring months. A field experiment was carried out to evaluate the effect of water availability during the pre‐flowering–flowering period on vegetative, reproductive and yield responses of olive trees grown in central Argentina. From the end of autumn to mid‐spring, four irrigation treatments were imposed to olive trees (Olea europaea, cv. Arbequina and Manzanilla) at 0, 25, 50 and 75% estimated crop evapotranspiration (ETc). Also, a control treatment was kept at 100% ETc for the entire year. For the first crop year evaluated, water deficit applied at early June, approximately 4 months prior to bloom, reduced the vegetative shoot growth and delayed the flowering time, resulting in shortening of the fruit maturation period and, ultimately, decreased fructification. Trees irrigated with high (75% of ETc) and full (100% of ETc) winter‐spring water supply presented significantly higher values of flower density, fruit density and final fruit yield which resulted in water productivity (kg fruits mm^?1 of irrigation/ha) enhancements of about 500% (cv. Arbequina) and 330% (cv. Manzanilla) with respect to those obtained from the corresponding unirrigated treatments. Differences between treatments in oil content and composition were primarily attributed to variations in fruit maturity. Differences in fatty acid composition were stronger in cv. Arbequina where a gradual increase in oleic acid content was registered in parallel to the increase in irrigation water supply. From a practical stand point, results obtained from most of the analysed parameters were quite similar for both T75 and T100 treatments. Thus, the possible convenience of irrigation at T75% ETc should be considered since it may warrant profitable olive production while saving a considerably quantity of irrigation water in the olive production area in central Argentina.     

Effects of PAR intensity and NaCl concentration on growth of <Emphasis Type="Italic">Salicornia europaea</Emphasis> plants as relevant to artificial ecological systems

Effects of variable levels of photosynthetically active radiation (PAR) and NaCl concentrations, typical of closed ecological life support systems, on growth of Salicornia europaea L. plants, CO₂ exchange, mineral composition, and the content of malondialdehyde (MDA) and photosynthetic pigments were investigated. The plants were grown for 25 days at different salinities of nutrient Knop solution (171, 342, and 513 mM NaCl) under two PAR levels (690 and 1150 μmol/(m² s)). At PAR of 690 μmol/(m² s), the plant productivity did not show significant changes at increasing salinities; at 1150 μmol/(m² s), the maximal productivity was observed at NaCl concentrations of 171 and 342 mM. The increase in NaCl concentration from 171 to 513 mM in the nutrient solution led to a substantial increase in the relative Na content in aboveground organs at PAR level of 1150 μmol/(m² s). The MDA content in aboveground organs by the end of the growth period was independent of PAR intensity. The content of photosynthetic pigments in the assimilatory tissue decreased with the increase in salinity from 342 to 513 mM NaCl at PAR level of 1150 μmol/(m² s) but not at the lower irradiance. The combination of 1150 μmol/(m² s) PAR intensity with the salinity as high as 342 mM NaCl was found to be the most effective for optimal productivity of S. europaea plants.     

Effects of fragmentation on forest structure and litter dynamics in Atlantic rainforest in Pernambuco, Brazil

This study evaluates the effects of fragmentation on the spatial and temporal dynamics of small litterfall production in Atlantic rainforest in Pernambuco State, Brazil. Litterfall was collected for 24 months at two 0.2 ha sites, located in the forest edge zone and the forest interior, within a rainforest patch of about 300 ha. Structural parameters of both forest sites were recorded. Litter was sorted into six fractions (foliage, twigs, buds/flowers, fruits/seeds, peduncles, rest), dried and weighed. The interior forest plot contained 314 live trees with a dbh 5 cm and a stand basal area of 41.8±8.7 m², whereas the forest edge contained 211 live trees and a stand basal area of 23.4±3.6 m². Total small litterfall was extraordinarily high and totalled 12.62±4.73 t ha⁻¹ yr⁻¹ in interior forest and 14.74±2.78 t ha⁻¹ yr⁻¹ in forest edge. High litterfall rates are probably due to a pronounced periodicity, edge effects alter litterfall strongly.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Leaf Gas Exchange and Water Relations in Polylepis tarapacana at Extreme Altitudes in the Bolivian Andes

Stress-induced restrictions to carbon balance, growth, and reproduction are the causes of tree-line formation at a global scale. We studied gas exchange and water relations of Polylepis tarapacana in the field, considering the possible effects of water stress limitations imposed on net photosynthetic rate (P _N). Daily courses of microclimatic variables, gas exchange, and leaf water potential were measured in both dry-cold and wet-warm seasons at an altitude of 4 300 m. Marked differences in environmental conditions between seasons resulted in differences for the dry-cold and wet-warm seasons in mean leaf water potentials (–1.67 and –1.02 MPa, respectively) and mean leaf conductances (33.5 and 58.9 mmol m^–2 s^–1, respectively), while differences in mean P _N (2.5 and 2.8 mol m^–2 s^–1, respectively) were not as evident. This may be related to limitations imposed by water deficit and lower photon flux densities during dry and wet seasons, respectively. Hence P. tarapacana has coupled its gas exchange characteristics to the extreme daily and seasonal variations in temperature and water availability of high elevations.