An integrated model for predicting maximum net photosynthetic rate of cocksfoot (Dactylis glomerata) leaves in silvopastoral systems

Net light-saturated photosynthetic rate (A_max) of field grown cocksfoot (Dactylis glomerata L.) leaves in a radiata pine (Pinus radiata D. Don) silvopastoral system (Canterbury, New Zealand) was measured at different times under severe shade (85–95 μmol m^–2 s^–1 photosynthetic photon flux density, PPFD) and in full sunlight (1900 μmol m^–2 s^–1 PPFD). The aim was to integrate individual functions for A_max against air temperature (2 to 37 oC), water status, expressed as pre-dawn leaf water potential (ψ_lp) (-0.01 to −1.6 MPa), herbage nitrogen (N) (1.5 to 5.9%), regrowth duration (20 to 60 days) and time under shade (1 to 180 min) into a multiplicative model. The highest A_max value obtained was 27.4 μmol CO₂ m^–2 s^–1 in non-limiting conditions with full sunlight. This value was defined as standardised dimensionless A_maxs = 1 for comparison of factor effects. The canopy temperature of the cocksfoot sward was up to 7.4 oC cooler than air temperature for plants under shade. Therefore, canopy temperature was used to predict A_max. The only interaction was between time under severe shade (5% of the open PPFD) and water stress (ψ_lp = −0.4 to −1.3 MPa) and this was included in the model. Validation of this model indicated 78% of the variation in A_max could be accounted for using these five factors by the addition of the interaction function. This model could be used to assist the prediction of pasture growth in silvopastoral systems through incorporation into a canopy photosynthesis model. This revised version was published online in June 2006 with corrections to the Cover Date.     

Growth of Festuca pallescens in Silvopastoral Systems in Patagonia, Part 1: Positive Balance between Competition and Facilitation

Silvopastoral systems may overyield compared to monocultures as a result of higher resource capture and/or facilitation effects of the trees on the pastures. Festuca pallescens (St. Ives) Parodi, a forage species of Patagonia, is vulnerable to water deficit, suggesting that it may benefit from the facilitative effects of trees. Radiation (PPFD), temperature, relative humidity (RH) and soil water distribution during the growing season were measured under different tree cover levels in ponderosa pine-based silvopastoral systems. PPFD decreased with increasing tree cover, and the decrease was lower in the positions under than between tree crowns. No significant differences were observed for temperature, RH nor for soil water content between treatments. Pre-dawn water potential of grasses during the growing season showed similar high values in all treatments. However, cumulative water stress was slightly lower in high tree cover treatments than in open situations. Leaf water potential of the grasses during the day was always lower in open grassland than in forested plots. The growth of the grasses showed no significant differences between treatments, but mean growth tended to decrease in the more densely planted treatments. We concluded that F. pallescens is a species that is biologically sound for use in silvopastoral systems, because the balance between facilitation and competition can be positive under the semiarid conditions of Patagonia.     

Photosynthetic responses to lightflecks ofFagus crenata seedlings grown in a gap and understory of a deciduous forest

Photosynthetic responses to a series of 1-min lightflecks (1,000μmol m⁻² s⁻¹) superimposed on a background with different duration (1, 5, and 10 min) and intensity (25 and 50μmol m⁻² s⁻¹) of low background photosynthetic photon flux density (PPFD) were measured in the leaves ofFagus crenata grown in a gap and understory of aFagus crenata forest in the Naeba Mountains. The two background PPFD intensities most frequently occurred in understory and gap sites respectively. The maximum net photosynthetic rate (P _Nmax) and maximum stomatal conductance (g _smax) were higher in the gap seedlings than in the understory seedlings. However, when the background PPFD was 25μmol m⁻²s⁻¹, the net photosynthetic rate (P ₂₅) and stomatal conductance (g _s25) were almost the same between the gap and understory. When the background PPFD duration was 1-min, the net photosynthetic rate (P _N ) at the end of each lightfleck increased progressively. When the background PPFD duration was 5- and 10-min, the increase inP _N at the end of each lightfleck was less. This indicates that background PPFD duration is important to photosynthetic responses to lightflecks. The higher ratios ofP ₂₅/P _Nmax andg _s25/g _smax in the understory seedlings indicate that the understory seedlings can maintain relatively lower levels of biochemical and stomatal limitations than the gap seedlings under low light conditions. The ratios ofP _N /P _Nmax at the end of each lightfleck (IS) and light utilization efficiency of single lightflecks (LUE _s) that showed the influence of lightflecks on carbon gain were higher in the understory seedlings than in the gap seedlings when the background PPFD was 25μmol m⁻² s⁻¹. This means that understory seedling are capable of utilizing fluctuating light more efficiently under low light conditions than the gap seedlings although the net carbon gain of single lightflecks (CG _s) in the understory seedlings was not higher than that in the gap seedlings. There were no significant differences inIS andLUE _s between understory seedlings at a background PPFD of 25μmol m⁻² s⁻¹ and gap seedlings at a background PPFD of 50μmol m⁻² s⁻¹. However,CG _s in gap seedlings was higher than in understory seedlings. These results provide more evidence thatF. crenata acclimate to a natural light environment in respect to relative induction state at low background PPFD and can capture the fluctuating light at the same efficiency in both the gap and understory seedlings under natural light environments. This study was funded by the research project, Evaluation of Total CO₂ Budget in Forest Ecosystems, coordinated by the Ministry of Agriculture, Forestry and Fisheries of Japan.     

Validation of a canopy photosynthesis model for cocksfoot pastures grown under different light regimes

Daily net canopy photosynthesis (P _n) was predicted for cocksfoot (Dactylis glomerata L.) canopies grown under different light regimes by integration of a leaf photosynthesis model developed for the light-saturated photosynthetic rate (P _max), photosynthetic efficiency (α) and the degree of curvature (θ) of the leaf light–response curve. When shade was the only limiting factor, the maximum P _n (P _nmax) was predicted to decrease approximately linearly from 33.4 g CO₂ m⁻² d⁻¹ to zero as photosynthetic photon flux density (PPFD) fell from full sunlight (1800 μmol m⁻² s⁻¹ PPFD) to 10% of this in a fluctuating light regime. It was also predicted that at 50% transmissivity P _nmax was higher for a continuous light regime (10.4 g CO₂ m⁻² d⁻¹) than for a fluctuating light regime with the same intensity (8.4 g CO₂ m⁻² d⁻¹). The canopy photosynthesis model was then used to predict dry matter (DM) production for cocksfoot field grown pastures under a diverse range of temperature, herbage nitrogen content and water status conditions in fluctuating light regimes. This prediction required inclusion of leaf area index and leaf canopy angle from field measurements. The model explained about 85% of the variation in observed cocksfoot DM production for a range from 6 to 118 kg DM ha⁻¹ d⁻¹. The proposed model improves understanding of pasture growth prediction through integration of relationships between shade limitations in fluctuating light regimes and other environmental factors that affect the canopy photosynthetic rate of cocksfoot pastures in silvopastoral systems.     

Modeling leaf maximum net photosynthetic rate of <Emphasis Type="Italic">Festuca pallescens</Emphasis>, the dominant perennial grass of Patagonian pine-based silvopastoral systems

The integrated relationship in a simple mechanistic model between the critical environmental factors controlling leaf photosynthesis of understory species would be a useful tool to optimize the management of the silvopastoral systems. Individual effect of leaf temperature, water stress and light environment over net maximum photosynthetic rate (Pmax) was evaluated on Festuca pallescens leaves grown in a silvopastoral system of two Pinus ponderosa canopy covers (350 and 500 trees ha⁻¹) and natural grassland. The aim was to integrate individual functions for Pmax against these environmental factors into a multiplicative model. We measured pre-dawn water potential (ψ _pd), leaf temperature and net photosynthetic rate (Pn), stomatal conductance (gs) and intercellular CO₂ concentration (Ci) as a function of photosynthetic photon flux density (PPFD). The highest Pmax under non-limiting conditions was 20.4 μmol CO₂ m⁻² s⁻¹ and was defined as standardized dimensionless Pmax _s  = 1 for comparison of environmental factors. The leaf temperature function showed an optimum range between 20.2 and 21.8°C where Pmax _s  = 1. Then, Pmax _s declined by an average 1 μmol CO₂ m⁻² s⁻¹ C⁻¹ from the optimum to 4.7 and 38.5°C. Pmax _s decreased at a rate of 9.49 μmol CO₂ m⁻² s⁻¹ MPa⁻¹ as water potential reaches −1.9 MPa and showed a lower slope as water potential decreased down to −4.3 MPa. The light environment was estimated from hemispherical photograph analysis. Pmax _s was 20% higher in leaves of open control plants than under the maximum tree canopy cover. The simple multiplicative model accounted for 0.82 of the variation in Pmax. Such a simple mechanistic model is the first step towards a more effective decision support tool.     

Shade acclimation in the forage grass Festuca Pallescens: biomass allocation and foliage orientation

Plants can acclimate to shade through different processes. In particular, they can modify their biomass allocation and the architecture in order to increase light interception. The objective of this study was to evaluate the shade acclimation capacity of Festuca pallescens (St. Ives) Parodi, as part of research concerning the use of this species in silvopastoral systems in Patagonia, Argentina. Biomass allocation was estimated from the leaf and root dry weights of plants growing in an open pasture and forested plots. Crown architecture of plants growing in the open and in two shade treatments was studied dividing each plant in three concentric cylinders, within which leaf angles and leaf area were measured. Light interception of plants in each treatment was estimated from the projected leaf areas and the relative amount of radiation reaching each location. Biomass allocation changed significantly in plants growing under shade conditions, increasing the proportion of leaves relative to the roots (Leaf Mass Fraction = 0.29 (SD: 0.12) and 0.40 (SD: 0.09) in plants in the open and under shade, respectively). Also, mean leaf inclination angles changed in plants growing under shade conditions, allowing an increase in light interception of approximately 35% compared to plants with the crown architecture typical of the open treatment. Previous studies have shown that F. pallescens does not change its photosynthetic response to light under shade conditions. Therefore, we conclude that the reported changes in biomass allocation and crown architecture, in addition to the increment in specific leaf area explain the relatively high shade tolerance of this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Changes in leaf water use after removal of leaf lower surface hairs on <Emphasis Type="Italic">Mallotus macrostachyus</Emphasis> (Euphorbiaceae) in a tropical secondary forest in Malaysia

Leaf hairs may assist in maintaining high leaf water use efficiency in tropical secondary forest tree species. We compared leaf temperature, transpiration, photosynthesis and water use efficiency between hairy and depilated leaves in Mallotus macrostachyus (Euphorbiaceae), to determine the role of leaf hair in leaf water use efficiency (WUE) in tropical degraded secondary forest in Malaysia. Measurements were made on five mature individuals growing in sun-exposed conditions and five in shaded conditions. The hair dry weight per unit leaf area was significantly greater in sun leaves than in shade leaves. The transpiration rate (Tr_max) of depilated leaves in sun-exposed conditions was slightly higher than in hairy leaves in both morning and afternoon measurements. In contrast, Tr_max in the shade leaves was almost identical in hairy and depilated leaves. Leaf stomatal conductance (g _s) in the morning showed almost the same value among leaf types and light conditions. In the afternoon, g _s slightly decreased from the morning values in both sun and shade conditions. In the morning, the leaf water use efficiency (A _max/Tr_max) in both conditions did not differ significantly between hairy and depilated leaves. However, in the afternoon, WUE in the depilated leaves was significantly lower than in hairy leaves in sun-exposed conditions. These observations suggest that leaf hairs in M. macrostachyus contribute to the high leaf water use efficiency in drought conditions, such as high vapor pressure deficit experienced at midday in degraded tropical secondary forests.     

Stomatal characteristics of Eucalyptus grandis clonal hybrids in response to water stress

This study describes the stomatal response occurring during water stress and subsequent recovery of three Eucalyptus grandis clonal hybrids. The aim was to investigate the degree to which stomatal conductance (g _s) and stomatal density differ between the clonal hybrids across seasons and in response to water stress. Plants from one E. grandis × E. camaldulensis (GC) and two E. grandis × E. urophylla (GU1 and GU2) clones were grown for 18 months in 80 l planting bags. Plants were subjected to three watering treatments: control (100% field capacity), chronic water stress (maintained at 15% of field capacity) and acute water stress (cyclic water stress, where water was withheld until leaf wilting point, and a subsequent period of recovery followed). Stomatal conductance was measured after 6, 12 and 18 months growth. At 12 months of age, the recovery of g _s 1, 2 and 7 d after rewatering (following acute water stress) was further investigated. The GC hybrid showed consistently higher g _s than the GU clones at each measurement period. Stomatal conductance was 24–66% higher during winter (after 12 months growth) than during summer. The recovery of stomatal conductance from acute water stress was more rapid in the GC clone than the GU clones. Chronic water stress was shown to decrease g _s in GU clones by up to 70%, but not in the GC clone. Water stress did not affect stomatal density or size. Remarkably, stomata were absent from the adaxial leaf surface of clone GU1 leaves, but not from the leaves of the other E. urophylla hybrid cross (GU2). Total biomass of the GC clone was significantly greater at 9 months growth, but after 18 months growth the GU1 clone had attained greater biomass accumulation (although not significantly). Measurement of g _s, transpiration, stomatal density and total biomass in the GU1 clone indicated stomatal sensitivity to water stress, a favourable trait during periods of drought. The differing growth strategies of the GU and GC clones could be partially explained by their differences in stomatal sensitivity in response to water stress.     

Short-term light and leaf photosynthetic dynamics affect estimates of daily understory photosynthesis in four tree species

Instantaneous measurements of photosynthesis are often implicitly or explicitly scaled to longer time frames to provide an understanding of plant performance in a given environment. For plants growing in a forest understory, results from photosynthetic light response curves in conjunction with diurnal light data are frequently extrapolated to daily photosynthesis (A(day)), ignoring dynamic photosynthetic responses to light. In this study, we evaluated the importance of two factors on A(day) estimates: dynamic physiological responses to photosynthetic photon flux density (PPFD); and time-resolution of the PPFD data used for modeling. We used a dynamic photosynthesis model to investigate how these factors interact with species-specific photosynthetic traits, forest type, and sky conditions to affect the accuracy of A(day) predictions. Increasing time-averaging of PPFD significantly increased the relative overestimation of A(day) similarly for all study species because of the nonlinear response of photosynthesis to PPFD (15% with 5-min PPFD means). Depending on the light environment characteristics and species-specific dynamic responses to PPFD, understory tree A(day) can be overestimated by 6-42% for the study species by ignoring these dynamics. Although these overestimates decrease under cloudy conditions where direct sunlight and consequently understory sunfleck radiation is reduced, they are still significant. Within a species, overestimation of A(day) as a result of ignoring dynamic responses was highly dependent on daily sunfleck PPFD and the frequency and irradiance of sunflecks. Overall, large overestimates of A(day) in understory trees may cause misleading inferences concerning species growth and competition in forest understories with < 2% full sunlight. We conclude that comparisons of A(day) among co-occurring understory species in deep shade will be enhanced by consideration of sunflecks by using high-resolution PPFD data and understanding the physiological responses to sunfleck variation.     

Photosynthetic response to variation in CO2 concentrations and temperature of four broad-leaved trees in Beijing region

Responses of the photosynthetic characteristics to variation in CO₂ concentration and temperature of Ginkgo biloba, Eucommia ulmoides, Magnolia denudata and Tilia japonica were measured during the peak growing season. The results show that the ambient CO₂ concentration could not meet the requirements for photosynthesis of these four species. The optimal temperatures for photosynthesis were lower than the average daytime air temperature. Hence, the photosynthesis of these four species was restricted by the low CO₂ concentration and high daytime air temperature at the time of measurement. Marked enhancements in the net photosynthetic rate were found in all four species when the CO₂ concentration was doubled. When the dependency on CO₂ and temperature were examined simultaneously, it was seen that for increased CO₂ concentrations there was a shift in the optimum temperature for M. denudata and T. japonica towards higher temperatures. Due to their independence on CO₂ concentrations, this trend could not be found in the G. biloba and E. ulmoides data sets. The stomatal conductance (G _s) was sensitive to a vapor pressure deficit (VPD) which in turn was sensitive to temperature. An increase in temperature would cause the VPD to increase and plants might be assumed to react by reducing their stomatal apertures. The effect on stomatal resistance would be most significant at high temperatures. The restriction to stomatal conductance for these four species would increase if CO₂ concentrations were elevated at the same temperature. __________ Translated from Journal of Agricultural University of Hebei, 2006, 29 (6): 39–43 [译自: 河北农业大学学报]     

Dry matter production,morphology and nutritive value of <Emphasis Type="Italic">Dactylis glomerata</Emphasis> growing under different light regimes

Plant growth, morphology and nutritive value under shade can differ between temperate grasses. Therefore, the aim of this study was to quantify the dry matter (DM) production, sward morphology, crude protein (CP%), organic matter digestibility (OMD) and macro-nutrient concentrations (P, K, Mg, Ca and S) in a grazed cocksfoot (Dactylis glomerata L.) pasture under 10-year-old Pinus radiata D. Don forest. Four levels of light intensity were compared: full sunlight (100% photosynthetic photon flux density-PPFD), open + wooden slats (∼43% PPFD), trees (∼58% PPFD) and tree + slats (∼24% PPFD). The mean total DM production was 8.2 t DM ha⁻¹ yr⁻¹ in the open and 3.8 t DM ha⁻¹ yr⁻¹ in the trees + slats treatment. The changes in cocksfoot leaf area index (LAI) were related to variations in morphological aspects of the sward such as canopy height and tiller population. CP% increased as PPFD declined with mean values of 18.6% in open and 22.5% in the trees + slats treatment. In contrast, the intensity of fluctuating shade had little effect on OMD with a mean value of 79 ± 3.2%. The mean annual macro-nutrient concentrations in leaves increased as the PPFD level declined mainly between the open and the trees + slats treatments. It therefore appears that heavily shaded dominant temperate pastures in silvopastoral systems limit animal production per hectare through lower DM production rates and per animal through reduced pre-grazing pasture mass of lower bulk density from the etiolated pasture.     

Effect of pruning on branch production and water relations in widely spaced ponderosa pines

This study dealt with the effects of pruning on branch and leaf area (F _a) production of ponderosa pines growing in silvopastoral systems in Patagonia. We hypothesized that pruning positively influences the number of branches per whorl and their basal area growth rate, changing F _a production. In addition, we studied some water relations in order to explain potential differences in branch growth rates between treatments. Two mathematical models were developed to estimate branch and total F _a. The averaged diameter at the third year of pruning was, for high-pruned trees 3.1 and 3.6 cm at the bottom and middle of the crown, against 4 and 4.4 cm for low-pruned trees. Pruning did not produce changes in the number of branches per whorl (approximately 7.6 branches per whorl). Water stress may be responsible of this lower branch growth in pruned trees. Water potential, stomatal conductance and transpiration were lower in high- than in low-pruned trees.     

生长抑制剂对大叶黄杨形态及光合作用的影响

目的]探究抑制剂对大叶黄杨生长的抑制作用以及对其叶片形态和光合作用的影响,为灌木绿篱的化学修剪提供技术指导。[方法]在北京林业大学林场苗圃采用3种生长抑制剂(多效唑(PP_(333))、三碘苯甲酸(TIBA)以及脱落酸(ABA))对密植成绿篱状的1年生大叶黄杨扦插苗进行叶面喷施,对其生长、叶片形态及光合作用等指标进行测定。[结果]3种抑制剂均有矮化植株、抑制新梢生长的作用,矮化效果最佳、抑制作用最强的为PP_(333),且高浓度PP_(333)对于高生长的抑制作用持效性较长。叶宽、叶厚以及叶面积在短期PP_(333)处理下高于对照。PP_(333)能提高大叶黄杨净光合速率,主要通过增加叶厚、气孔导度、叶肉导度及叶绿素含量来实现,并且PP_(333)使蒸腾速率提高的同时降低了水分利用效率。TIBA有显著减小叶长、叶宽的作用,但能使叶厚增加,且随浓度的增加作用效果增强,主要通过减小叶面积,降低叶肉导度、气孔导度、胞间CO_2浓度及叶绿素含量来降低净光合速率。ABA能增加叶面积(10 mg·L~(-1)处理除外)、叶长以及减小叶片厚度,通过抑制气孔导度、叶肉导度、叶绿素含量降低净光合速率。TI-BA及ABA均通过降低蒸腾速率,使水分利用效率提高。[结论]3种抑制剂均有矮化植株、抑制新梢生长的作用,并且影响叶片发育,进而影响叶片光合作用,且800 mg·L~(-1)的多效唑对大叶黄杨具有较好的正向作用。     

Mesophyll conductance to CO2 and leaf morphological characteristics under drought stress during Quercus ilex L. resprouting

? Quercus ilex L., the dominant species in Mediterranean forests and one with a great capacity for resprouting after disturbances, is threatened by the expected increase in fire frequency and drought associated with climate change.

? The aim of this study was to determine the contribution of photosynthesis limitants, especially mesophyll conductance (g _mes) during this species’ resprouting and under summer drought.

? Resprouts showed 5.3-fold increased g _mes and 3.8-fold increased stomatal conductance (g _s) at mid-day with respect to leaves of undisturbed individuals. With increased drought, structural changes (decreased density and increased thickness) in resprouts contributed to the observed higher photosynthesis and increased g _mes. However, g _mes only partially depended on leaf structure, and was also under physiological control. Resprouts also showed lower non-stomatal limitations (around 50% higher carboxylation velocity (V _c,max) and capacity for ribulose-1,5-bisphosphate regeneration (J _max)). A significant contribution of g _mes to leaf carbon isotope discrimination values was observed.

? g _mes exhibits a dominant role in photosynthesis limitation in Q. ilex and is regulated by factors other than morphology. During resprouting after disturbances, greater capacity to withstand drought, as evidenced by higher g _mes, g _s and lower non-stomatal limitants, enables increased photosynthesis and rapid growth.

     

Physiological responses of Eucalyptus nitens × nitens under experimentally imposed water stress

This study reports on how a cold- and frost-tolerant clone of Eucalyptus nitens × nitens responds to drought stress. The aim was to identify physiological traits that contribute to drought acclimation. Macropropagated saplings were grown in a climate-controlled greenhouse in pots filled with coarse river sand supplied with a slow-release fertiliser. One group of plants was kept regularly watered (control), and another group was subjected to four cycles of water stress (drought) whereby water was withheld for periods lasting 6, 10, 10 and 14 d, with 4 d of regular watering (recovery) inbetween. A drought cycle was terminated once saplings showed signs of wilting. Daily responses in stomatal conductance (g _s) were similar between control and drought treatments, except on the day of termination of the cycle, when g _s was significantly depressed in droughted plants. During the fourth and most severe drought cycle, there was a physiological adaptation to water stress because g _s was similar between control and droughted plants. Stomatal conductance was significantly positively correlated with volumetric soil moisture content in the drought treatment, but not in the control treatment. Electron transport capacity (J _max) increased during each drought cycle, and the increase was significant during the fourth cycle. Other parameters derived from A/c _i response curves were similar between the treatments. Under experimentally imposed water stress, E. nitens × nitens reduced leaf area, increased assimilate rate per unit leaf area, and maintained high stomatal conductance until leaves wilted. After 46 d droughted plants had accumulated half the biomass of control plants. Therefore, a cold- and frost-tolerant clone of E. nitens × nitens may be tolerant to drought stress but at a reduced growth rate because of reduced leaf area.     

Evaluation of plantation and early development of five alternatives to ponderosa pine in silvopastoral systems in northwest Patagonia,Argentina

Previous studies of afforestation in Patagonia indicate that 30–50 % tree cover produces positive effects on the pasture. This coverage level is achieved by applying pruning and strong thinning to reduce the volume of timber production per area unit. From an economic standpoint, in order to not reduce the income level of the system, it is necessary to find tree species that could replace ponderosa pine (Pinus ponderosa) and maximize income per volume unit of wood. We evaluated the feasibility of implementation of five broadleaved tree species (native and exotic) with higher intrinsic wood quality than ponderosa pine. We tested the influence of tree cover generated by a framework of silvopastoral plantation on the increase in survival, regrowth and absolute increase in height, and compared ecophysiological variables (net photosynthetic activity, stomatal conductance, intrinsic water use efficiency) of different species to the status of a traditional plantation without tree cover. Additionally values of air temperature and relative humidity were registered under both cover conditions. Preliminary results support the conclusion that three of these five species could be considered as alternatives to ponderosa pine species for establishing silvopastoral systems with higher timber value. Our Based on the results, the use of tree cover generated under the current plantations of P. ponderosa could increase the success of the establishment of these species.     

Carbon Isotope Discrimination, Gas Exchange and Stem Growth of Four Euramerican Hybrid Poplars under Different Watering Regimes

Seasonal changes in carbon isotope discrimination (Δ) and gas exchange traits were assessed in four Populus×euramericana clones differing in growth potential. Measurements were made during the second year after establishment in the field under two watering regimes, which were defined by the time-span between flood irrigations, hence resulting in different dry-down cycles: high irrigation (conservative schedule currently applied in the Ebro Valley, Spain) and low irrigation (equivalent to about a one-fourth reduction in water inputs). Net CO₂ assimilation rate (A), stomatal conductance (g_s), intrinsic water-use efficiency (A/g_s) and other related photosynthetic traits (leaf nitrogen concentration, leaf greenness and leaf mass per area) were measured prior to watering, and Δ was analysed in water-soluble leaf extracts (Δ_s) and bulk leaves (Δ_l). Stem growth was monitored over 3 years starting at the year of establishment (1998). Data were subjected to a repeated measures ANOVA over time for a randomised block split-plot design across watering regimes. Significant differences between watering regimes were detected using a long-term estimate of photosynthetic performance such as Δ_l, in agreement with changes in soil water status and evapotranspirative demand. However, the lack of significant genotype×watering regime interactions for gas exchange traits and Δ_s suggested that water shortage imposed by low irrigation was not sufficient to reveal physiological adaptations to drought. In this regard, the reduction in water inputs brought about by low irrigation did not reduce tree growth for any of the clones, suggesting that the current irrigation scheme employed in the region is superfluous to the water consumption needs of poplars. Genotypic variation was detected in gas exchange traits, Δ_s, Δ_l and stem growth under both watering treatments. Significant correlations with stem volume for Δ_s (r = −0.60, p<0.05) and A (r = + 0.61, p<0.05) suggested that growth was improved by higher water-use efficiency (the ratio of carbon fixed to water lost, as inferred by Δ_s) due to variation in A rather than in g_s. This observation corroborated the expectation derived from current theories that a lower Δ is related to higher stem volume, as a result of changes in net CO₂ assimilation rates.     

Spatial and seasonal variations in photosynthetic properties within a beech (Fagus crenata Blume) crown

The light response curve and the intercellular CO₂ concentration response curve of CO₂ assimilation rate were investigated together with the light conditions at the four different heights within the beech crown from 1995 to 1997 on Mt. Fuji in Japan. On the seasonal fluctuation, the CO₂ assimilation rate at light saturated condition increased rapidly in May, and attained to the maximum between the end of June and July, thereafter, slightly decreased until the middle of August and rapidly decreased in September and October. The daily sum of photosynthetic photon flux density attenuated with deeping within the crown, and particularly, the relative value on 2nd position dropped to only 30%. TheA _max decreased from 10 to 5μmol m⁻² s⁻¹, approximately, with deeping within the crown. The light saturation point, quantum yield, light compensation point and dark respiration rate also varied with deeping. These results suggest that the photosynthetic properties vary gradually from sun to shade leaves along the light attenuation within a beech crown. At light saturated condition, the stomatal conductance and mesophyll conductance were strongly correlated withA _max among the four different heights (r > 0.96, respectively). TheC _i/C _a ratio was around 0.8, and there were no remarkable differences among the four different heights. These results suggest that the vertical gradient ofA _max depends on the variation of mesophyll conductance. The stomatal conductance may be also one of the major factor in the vertical gradient ofA _max. However the intercellular CO₂ concentration doesn’t influence the vertical gradient ofA _max within the crown. This work is supported by the Sasagawa Scientific Research Grant from The Japan Science Society and Grant-in-Aid for Scientific Research (C).     

Silvopastoral systems in northwestern Patagonia. I: growth and photosynthesis of Stipa speciosa under different levels of Pinus ponderosa cover

Combination of native pastures with fast-growing tree species (mainly Pinus spp.), may be an interesting economic and environmental alternative for small and medium land-owners in Patagonia,Argentina. Pasture productivity is usually the single most important factor affecting livestock carrying capacity. Therefore the prediction of the effects of tree cover on understory herbaceous production is of great importance for management. This paper reports the growth response of Stipa speciosa Trin. et Rupr. (tillering, tiller height, number of green leaves per tiller, and relative growth) to Pinus ponderosa Doug. (Laws) canopy cover (0 to 100%). Grasses did not grow when crown cover was >70%. Tiller production and relative growth were negatively correlated with tree cover; tiller height was larger under high tree cover. There was no relationship between leaf production and tree canopy cover. Grass leaves maintained net photosynthetic rates, at similar light levels, at a wide range of plant water potentials (from saturation to at least –2.4 MPa). Decrease in photosynthesis occurred at very low plant water potentials (–4.3 MPa). No differences in photosynthetic rate at similar radiation levels were foundbe tween plants growing in different light environments (determined by tree cover) in the field. Stipa speciosa can maintain positivenet CO₂ assimilation and growth under moderate shading and severewater stress conditions. For this reason, it can be used in silvopastoral systems, with a proper light management.This revised version was published online in November 2005 with corrections to the Cover Date.     

Diurnal Course of Gas Exchange and Water Use Efficiency of Platycladus orientalis in the Semiarid Region of the Loess Plateau

The daily gas exchange, stomatal conductance, and water use efficiency of Platycladus orientalis in the test field of Fangshan county of Shanxi Province in the semiarid region of the Loess Plateau were measured with portable Li-6200 gas analysis system in natural conditions. The results showed that the diurnal course of net photosynthetic rate displayed a two-peak pattern, that of stomatal conductance displayed a hollow pattern, and that of transpiration rate displayed a signal-peak pattern. Water use efficiency culminated in the early morning. On the basis of two criteria of stomatal limitation of photosynthesis suggested by Farquhar and Sharkey, the predominant limiting factor of photosynthesis was the stomatal conductance of stomatal limitation in the morning (10:00–12:00). However, the midday depression of photosynthesis at noon (12:00–14:00) and the decrease in photosynthesis in the afternoon (16:00–18:00) were the results of nonstomatal limitation, such as low carboxylation capacity of the leaf mesophyll.