Distribution of spore-positive and spore-negative nodules in stands ofAlnus glutinosa andAlnus incana in Finland

Summary The distribution of spore positive (Sp⁺) and spore negative (Sp⁻) nodules on the two native alder species (A. incana andA. glutinosa) in Finland was investigated. Nodules were collected throughout the country from different ecosystems (forests, swamps, lake- sea- and riversides, old pastures and fields as well as from alder plantations). OnA. incana Sp⁺ nodules predominated, whereas onA. glutinosa the vast majority of the nodules were of the Sp⁻ type. Sp⁺ nodules onA. glutinosa were found only at sites where the two alder species grew close together. This distribution pattern indicates an association of nodule type with alder species, the reasons for which are discussed. Indications of saprophytic growth in the Sp⁻ strain were also found.     

Day-to-day variation in nitrogenase activity of Alnus incana explained by weather variables: a multivariate time series analysis

A modelling system is described that indicates the extent to which day-to-day variations in nitrogenase activity in young Alnus incana (L.) Moench, grown in defined conditions in the field, may be affected by weather conditions both during and prior to the day of measurement. Nitrogenase activity (acetylene reduction activity, ARA) was measured weekly on intact field-grown grey alder (A. incana) plants, 0.15–0.42 m tall at planting, nodulated with Frankia. The measurements were done at noon on two groups of plants in 1987 and on two other groups in 1988. Each group was made up of five or six plants. Seven weather variables: daily sunshine hours, daily mean, maximum and minimum air temperature, daily mean and 1300 h relative humidity, and daily rainfall were used. The relation between log(ARA/leaf area) and the weather variables were analysed using a PLS model (partial least squares projection to latent structures). The advantage of PLS is that it can handle x-variables that are correlated. Data from 1987 were chosen as a training set. Multivariate PLS time series analysis was made by adding, in a stepwise manner, the weather data up to 5 d before the day of measurement. This procedure gave six models with n * 7 x-variables (n= 1–6). With the models from the time series analysis of 1987 data, true predictions of ARA per leaf area were made from weather data 1988 (test set 1) and from ‘early-season’ weather data from 1987 and 1988 (test set 2). The variation in ARA/leaf area could be predicted from the weather conditions. The predictions of the two test sets improved when the weather conditions one and two days before the day of measurements were added to the model. The further addition of weather data from 3 to 5 d before the day of measurement did not improve the model. The good predictions of ARA/leaf area show that the alders responded to the variable weather conditions in the same way in 1988 as in 1987, despite the ten-fold difference in size (leaf area) at the end of the growing season. Among the weather variables, air temperature and the daily sunshine hours were positively correlated to ARA, while relative air humidity and rainfall were negatively correlated to ARA. The daily minimum temperature and rainfall appeared to have least impact on ARA. By use of PLS, we could extract information out of a data set containing highly correlated x-variables, information that is non-accessible with conventional statistical tools such as multiple regression. When making measurements of nitrogenase activities under field conditions, we propose that attention should be paid to the weather conditions on the days preceding the day of measurement. The day-to-day variation in nitrogenase activity is discussed with reference to known effects of stress factors under controlled conditions.     

Alder and lupine enhance nitrogen cycling in a degraded forest soil in Northern Sweden

Positive effects of legumes and actinorhizal plants on N-poor soils have been observed in many studies but few have been done at high latitudes, which was the location of our study. We measured N₂ fixation and several indices of soil N at a site near the Arctic Circle in northern Sweden. More than 20 years ago lupine (Lupinus nootkatensis Donn) and gray alder (Alnus incana L. Moench) were planted on this degraded forest site. We measured total soil N, net N mineralization and nitrification with a buried bag technique, and fluxes of NH⁺ ₄ and NO^– ₃ as collected on ion exchange membranes. We also estimated N₂ fixation activity of the N₂-fixing plants by the natural abundance of ¹⁵N of leaves with Betula pendula Roth. as reference species. Foliar nitrogen in the N₂-fixing plants was almost totally derived from N₂ fixation. Plots containing N₂-fixing species generally had significantly higher soil N and N availability than a control plot without N₂-fixing plants. Taken together, all measurements indicated that N₂-fixing plants can be used to effectively improve soil fertility at high latitudes in northern Sweden.     

Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro

Cultures in vitro of Betula pendula Roth were subjected to light of different spectral qualities. Photosynthetic capacity was highest when the plantlets were exposed to blue light (max recorded photosynthesis, 82 mol CO₂ dm^–2 h^–1) and lowest when irradiated with light high in red and/or far-red wave lengths (max recorded photosynthesis, 40 mol CO₂ dm^–2 h^–1). Highest chlorophyll content (2.2 mg dm^–2 leaf area) was found in cultures irradiated with blue light, which also enhanced the leaf area. Morphometric analysis of light micrographs showed that the epidermal cell areas were largest in plantlets subjected to blue light and smallest in those subjected to red light. Morphometric analysis of electron micrographs of palisade cells, showed that the functional chloroplast area was largest in chloroplasts of leaves subjected to blue light and smallest in those exposed to red light. We suggest that light quality affects photosynthesis both through effects on the composition of the photosynthetic apparatus and on translocation of carbohydrates from chloroplasts.     

In vitro propagation of mature trees of Alnus incana (L.) Moench.

Mature trees of European grey alder (Alnus incana) were micropropagated on a modified MS medium containing 2.5 M BA, 6.2 mM (500 mg l^-1) NH₄NO₃ and 1.5% glucose. Prior to in vitro culture, mature scions were multiplied through grafting and cutting techniques. Shoot tips from cuttings were established in vitro. After six months of culture, shoots were rooted either in vitro or in vivo and plantlets were transferred to greenhouse conditions.     

Improvement of growth conditions and gas exchange of<Emphasis Type="Italic"> Fagus sylvatica</Emphasis> L. seedlings planted below a recently thinned<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> L. stand

Abstract Gas exchange and growth of beech seedlings planted in the understory of a recently thinned pinewood were recorded for 2 years. Relative irradiance was assessed by hemispherical photographs taken just after the thinning. Predawn water potential (_pd), daily gas exchange and chlorophyll fluorescence were measured several times during the two growing seasons. Maximum values of photosynthesis (A _max) and stomatal conductance to water vapour (g _wvmax) were established from daily data. Maximum quantum efficiency of PS II was recorded at dawn by taking the variable to maximum chlorophyll fluorescence ratio on dark adapted leaves (F _v/F _m). In the middle of each summer, leaf nitrogen content and leaf mass per area were evaluated, and height growth and basal area increment were recorded at the end of the season. The thinning treatment removed half the trees and generated around 10% more available relative irradiance (GLF). This was followed by an increase in net photosynthesis at saturating PPFD (A _sat) and in maximum stomatal conductance to water vapour (g _wvmax). Moreover, specific leaf mass (SLM) and mass based nitrogen content (N_m) showed higher values for seedlings in the thinned stand. In both years, a positive relationship was established between the area based nitrogen content (N_a) and maximum net photosynthesis (A _max). In 1998, a year with a dry summer, seedlings suffered a significant drop in daily A _max irrespective of the thinning regime. This was a response to an increase in stomatal limitation to net photosynthesis, g _wvmax reaching the lowest value on dates with the highest drought. A lack of decrease of Fv/Fm confirmed the absence of significant non-stomatal limitation to A as a consequence of photoinhibition after opening the pinewood. A higher maximum quantum efficiency of open PS II centres (Fv/Fm) was registered in seedlings in the thinned stand. The significance of the differences between the treatments was stronger in the second year after thinning. In 1999, a year with frequent summer storms, water availability increased for seedlings growing under the thinned pinewood. Overall, the reduced pine overstory had a positive effect on physiological responses of beech seedlings, which was translated into improved seedling growth.     

C4 photosynthetic carbon metabolism in the leaves of aromatic tropical grasses — I. Leaf anatomy,CO2 compensation point and CO2 assimilation

A few species of Cymbopogon and Vetiveria are potentially important tropical grasses producing essential oils. In the present study, we report on the leaf anatomy and photosynthetic carbon assimilation in five species of Cymbopogon and Vetiveria zizanioides. Kranz-type leaf anatomy with a centrifugal distribution of chloroplasts and exclusive localization of starch in the bundle sheath cells were common among the test plants. Besides the Kranz leaf anatomy, these grasses displayed other typical C₄ characteristics including a low (0–5 µl/l) CO₂ compensation point, lack of light saturation of CO₂ uptake at high photon flux densities, high temperature (35°C) optimum of net photosynthesis, high rates of net photosynthesis (55–67 mg CO₂ dm^-2 leaf area h^-1), little or no response of net photosynthesis to atmospheric levels of O₂ and high leaf ¹³C/¹²C ratios. The biochemical studies with ¹⁴CO₂ indicated that the leaves of the above plant species synthesize predominantly malate during short term (5 s) photosynthesis. In pulse-chase experiments it was shown that the synthesis of 3-phosphoglycerate proceeds at the expense of malate, the major first formed product of photosynthesis in these plant species.     

Study of the contribution of the shoot and/or root ofAlnus sp. in the compatibility between the host and a Sp+ Frankia strain using a grafting technique

Two alder species,Alnus glutinosa (L.) Gaertn. andAlnus incana (L) Moench, were inoculated with a Sp⁺ Frankia homogenate obtained fromA. incana root nodules. This inoculum formed effective nodules on the original host plant and ineffective nodules onA. glutinosa. Grafts between the two alder species were made to determine which part of the plant is involved in this phenomenon. The results obtained indicate that the compatibility between Alnus andFrankia is restricted to the root system.     

Acclimation to light in seedlings of <Emphasis Type="Italic">Quercus petraea</Emphasis> (Mattuschka) Liebl. and <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd. planted along a forest-edge gradient

Photosynthetic acclimation of two co-occurring deciduous oaks (Quercus petraea and Quercus pyrenaica) to a natural light gradient was studied during one growing season. In the spring of 2003, 90 seedlings per species were planted along a transect resulting from a dense Pinus sylvestris stand, an adjacent thinned area and a 10-m-wide firebreak (16.5–60.9% Global Site Factor (GSF)). In two dates of the following summer, we measured leaf gas exchange, carboxylation efficiency (CE), chlorophyll and nitrogen content, light–response curves of chlorophyll a fluorescence parameters, and leaf mass per area (LMA). Summer was mild, as evidenced by leaf predawn water potential (Ψ_pd), which reduced the interactive effect of water stress on the response of seedlings to light. Q. pyrenaica had higher LMA, CE, stomatal conductance (g _{s max}) and photosynthesis per unit area than Q. petraea at all growth irradiances. , LMA, g _{s max} and electron transport rate (ETR) all increased with light availability (GSF) in a similar fashion in both species. Light had also a clear effect on the organization of Photosystem II (PS II), as deduced by chlorophyll a fluorescence curves. Chlorophyll concentration (Chl_m) decreased with increasing light availability in Q. pyrenaica but it did not in Q. petraea. Seedlings of Q. petraea acclimated to higher irradiances showed a greater non-photochemical quenching (NPQ) than those of Q. pyrenaica. This suggests a higher susceptibility to high light in Q. petraea, which would be consistent with a better adaptation to shade, inferred from the lower LMA or the lower rate of photosynthesis.     

Functional Diversity of Culturable Bacterial Communities in the Rhizosphere in Relation to Fine-root and Soil Parameters in Alder Stands on Forest, Abandoned Agricultural, and Oil-shale Mining Areas

Grey alder (Alnus incana) and black alder (Alnus glutinosa) stands on forest land, abandoned agricultural, and reclaimed oil-shale mining areas were investigated with the aim of analysing the functional diversity and activity of microbial communities in the soil–root interface and in the bulk soil in relation to fine-root parameters, alder species, and soil type. Biolog Ecoplates were used to determine community-level physiological profiles (CLPP) of culturable bacteria in soil–root interface and bulk soil samples. CLPP were summarized as AWCD (average well color development, OD 48 h⁻¹) and by Shannon diversity index, which varied between 4.3 and 4.6 for soil–root interface. The soil–root interface/bulk soil ratio of AWCD was estimated. Substrate-induced respiration (SIR) and basal respiration (BAS) of bulk soil samples were measured and metabolic quotient (Q = BAS/SIR) was calculated. SIR and Q varied from 0.24 to 2.89 mg C g⁻¹ and from 0.12 to 0.51, respectively. Short-root morphological studies were carried out by WinRHIZO^TM Pro 2003b; mean specific root area (SRA) varied for grey alder and black alder from 69 to 103 and from 54 to 155 m² kg⁻¹, respectively. The greatest differences between AWCD values of culturable bacterial communities in soil–root interface and bulk soil were found for the young alder stands on oil-shale mining spoil and on abandoned agricultural land. Soil–root interface/bulk soil AWCD ratio, ratio for Shannon diversity indices, and SRA were positively correlated. Foliar assimilation efficiency (FOE) was negatively correlated with soil–root interface/bulk soil AWCD ratio. The impact of soil and alder species on short-root morphology was significant; short-root tip volume and mass were greater for black alder than grey alder. For the investigated microbiological characteristics, no alder-species-related differences were revealed.     

Effects of an Experimental Increase of Temperature and Drought on the Photosynthetic Performance of Two Ericaceous Shrub Species Along a North–South European Gradient

Plant ecophysiological changes in response to climatic change may be different in northern and southern European countries because different abiotic factors constrain plant physiological activity. We studied the effects of experimental warming and drought on the photosynthetic performance of two ericaceous shrubs (Erica multiflora and Calluna vulgaris) along a European gradient of temperature and precipitation (UK, Denmark, The Netherlands, and Spain). At each site, a passive warming treatment was applied during the night throughout the whole year, whereas the drought treatment excluded rain events over 6–10 weeks during the growing season. We measured leaf gas exchange, chlorophyll a fluorescence, and leaf carbon isotope ratio (¹³C) during the growing seasons of 1999 and 2000. Leaf net photosynthetic rates clearly followed a gradient from northern to southern countries in agreement with the geographical gradient in water availability. Accordingly, there was a strong correlation between net photosynthetic rates and the accumulated rainfall over the growing season. Droughted plants showed lower leaf gas exchange rates than control plants in the four sites. Interestingly, although leaf photosynthetic rates decreased along the precipitation gradient and in response to drought treatment, droughted plants were able to maintain higher leaf photosynthetic rates than control plants in relation to the accumulated rainfall over the months previous to the measurements. Droughted plants also showed higher values of potential photochemical efficiency (F _v/F _m) in relation to controls, mainly at midday. The warming treatment did not affect significantly any of the studied instantaneous ecophysiological variables..     

Effect of the D genome and of selection on photosynthesis in wheat

Summary Photosynthesis and transpiration in wheats and in their progenitors were analyzed in relation to their genome, ploidy and selection. The values of these parameters markedly depend on a specific effect of the D genome and on leaf enlargement in the course of evolution in wheats. Leaf enlargement has had a marked effect on photosynthesis in the genotypes that are devoid of the D genome; in addition, their photosynthetic capacity is greater in forms with lower leaf area. The increase in the mesophyll resistance r_m to CO₂ transfer is in relation to the increase in leaf area and is mainly responsible for the decrease in photosynthesis rate.Owing to its stomatal regulation, Triticum aestivum L. is characterized by good water use efficiency in spite of its large leaves and of its low net photosynthesis. On the basis of the photosynthesis rate, the large leaf factor does not appear to be a good selection criterion for the Triticum durum genotypes that are devoid of the D genome.     

Nitrogen in shoot litter,root litter and root exudates from nitrogen-fixingAlnus incana

Summary A pot experiment withAlnus incana (L.) Moench growing in sand was set up to compare the amounts of nitrogen released from plants shoot litter with that released below ground as root litter and/or root exudation. No nitrogen fixation by free-living microorganisms was found in the sand and the increased nitrogen content of the plant + soil system was therefore due to nitrogen fixation byFrankia in the alder root-nodules. Most of the nitrogen released from the plants was in the nitrogen-rich leaf and other shoot litter. Only small amounts of nitrogen were found in the drainage water from the pots and were recorded as increased nitrogen content of the sand.     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.     

Nitrogen and carbon fixation byAnabaena sp. isolated from a rice paddy and grown under P and light limitations

Anabaena sp., isolated from a rice paddy, was investigated for its nitrogen fixation as measured by acetylene reduction activity (ARA) in P-limited continuous and light-limited semi-continuous cultures. Growth rate (μ) under P limitation was a function of cell P content (q _p). Both the photosynthetic capacity (P_max) and photosynthetic efficiency (α) increased with μ when expressed per cell, but not per unit chla. The ARA of steady-state cells under P limitation increased with μ and was linearly related to C-fixation rate. This was apparently a consequence of the control of C-fixation by P limitation. In light-limited cells, steady state ARA, both at the culture light intensity and in the dark, increased asymptotically with μ, but the activity in the dark was only about 51% of that in the light. When the light level of steady-state cells grown at a high in intensity was switched to a low level, ARA decreased exponentially with time. Dark ARA activity also showed a similar decline, but at much lower levels. Thus, ARA depended not only on light history, but also immediate photosynthesis. Steady-state ARA at the ambient intensity or in the dark showed a strong correlation with¹⁴C-fixation rate. ARA of light-limited cells showed the same light-saturation characteristics as their¹⁴C-fixation, with the same initial saturation intensity,I _k. The ratios of P_max to the maximum ARA (ARA_max), and α to the slope of ARA (α_ara) were identical. A comparison of gross to net photosynthesis and N₂ fixation suggested that there was little leakage or excretion of fixed C or N.     

Leaf and canopy photosynthetic CO2 uptake of a stand of Echinochloa polystachya on the Central Amazon floodplain

The C₄ grass Echinochloa polystachya, which forms dense and extensive monotypic stands on the Varzea floodplains of the Amazon region, provides the most productive natural higher plant communities known. The seasonal cycle of growth of this plant is closely linked to the annual rise and fall of water level over the floodplain surface. Diurnal cycles of leaf photosynthesis and transpiration were measured at monthly intervals, in parallel with measurements of leaf area index, canopy light interception and biomass. By artificial manipulation of the light flux incident on leaves in the field light-response curves of photosynthesis at the top and near to the base of the canopy were generated. Fitted light-response curves of CO₂ uptake were combined with information of leaf area index, incident light and light penetration of the canopy to estimate canopy rates of photosynthesis. Throughout the period in which the floodplains were submerged photosynthetic rates of CO₂ uptake (A) for the emergent leaves were high with a mean of c. 30 mol m^-2 s^-1 at mid-day and occasional values of 40 mol m^-2 s^-1. During the brief dry phase, when the floodplain surface is uncovered, there was a significant depression of A, with mid-day mean values of c. 17 mol m^-2 s^-1. This corresponded with a c. 50% decrease in stomatal conductance, and a c. 35% depression in the ratio of the leaf inter-cellular to external CO₂ concentration (c _i/c _a). During the dry phase, a midday depression of rates of CO₂ assimilation was observed. The lowest leaf area index (F) was c. 2 in November–December, when the flood plain was dry, and again in May, when the rising floodwaters were submerging leaves faster than they were replaced. The maximum F of c. 5 was in August when the floodwaters were receding rapidly. Canopy light interception efficiency varied from 0.90 to 0.98. Calculated rates of canopy photosynthesis exceeded 18 mol C m^-2 mo^-1 throughout the period of flooding, with a peak of 37 mol C m^-2 mo^-1 in August, but declined to 13 mol C m^-2 mo^-1 in November during the dry phase. Estimated uptake of carbon by the canopy from the atmosphere, over 12 months, was 3.57 kg C m^-2. This was insufficient to account for the 3.99 kg C m^-2 of net primary production, measured simultaneously by destructive harvesting. It is postulated that this discrepancy might be accounted for by internal diffusion of CO₂ from the CO₂-rich waters and sediments via the roots and stems to the sites of assimilation in the leaves.     

Canopy structure and vertical patterns of photosynthesis and related leaf traits in a deciduous forest

Canopy structure and light interception were measured in an 18-m tall, closed canopy deciduous forest of sugar maple (Acer saccharum) in southwestern Wisconsin, USA, and related to leaf structural characteristics, N content, and leaf photosynthetic capacity. Light attenuation in the forest occurred primarily in the upper and middle portions of the canopy. Forest stand leaf area index (LAI) and its distribution with respect to canopy height were estimated from canopy transmittance values independently verified with a combined leaf litterfall and point-intersect method. Leaf mass, N and A _max per unit area (LMA, N/area and A _max/area, respectively) all decreased continuously by over two-fold from the upper to lower canopy, and these traits were strongly correlated with cumulative leaf area above the leaf position in the canopy. In contrast, neither N concentration nor A _max per unit mass varied significantly in relation to the vertical canopy gradient. Since leaf N concentration showed no consistent pattern with respect to canopy position, the observed vertical pattern in N/area is a direct consequence of vertical variation of LMA. N/area and LMA were strongly correlated with A _max/area among different canopy positions (r²=0.81 and r²=0.66, respectively), indicating that vertical variation in area-based photosynthetic capacity can also be attributed to variation in LMA. A model of whole-canopy photosynthesis was used to show that observed or hypothetical canopy mass distributions toward higher LMA (and hence higher N/area) in the upper portions of the canopy tended to increase integrated daily canopy photosynthesis over other LMA distribution patterns. Empirical relationships between leaf and canopy-level characteristics may help resolve problems associated with scaling gas exchange measurements made at the leaf level to the individual tree crown and forest canopy-level.     

Gas exchange and photosynthesis of Eucalyptus camaldulensis seedlings inoculated with different ectomycorrhizal symbionts

Eucalyptus camaldulensis Dehnh. seedlings inoculated with Pisolithus tinctorius (Pers.) Coker & Couch and Thelephora terrestris Ehrl. per Fr. were grown in well watered soil (_s –0.03 MPa) or subjected to a long-term soil water stress of up to –1.0 MPa over 13-week period in a glasshouse. After 13 weeks, all seedling containers were watered to field capacity and then water was withheld from the E. camaldulensis seedlings to induce a short-term drought. Diurnal measurements of seedling photosynthesis rate (A), leaf stomatal conductance (g) and leaf water potential (_p) were completed before, during, and after the short term drought. Although they were growing in an equal soil volume, photosynthesis rate (A), leaf stomatal conductance and leaf water potential (_p) of larger seedlings with P. tinctorius ectomycorrhizae were similar to those of smaller seedlings colonized with T. terrestris during the short-term drought period. Seedlings inoculated with Pisolithus tinctorius maintained higher photosynthesis rates over the course of the short-term drought. Thus, P. tinctorius ectomycorrhizae appear to be more efficient than those of T. terrestris in assisting seedlings to maintain gas exchange and photosynthesis under limited soil moisture conditions.     

Effects of leaf position and nitrogen supply on the expansion of leaves of field grown sunflower (Helianthus annuus L.)

Leaf growth responses to N supply and leaf position were studied using widely-spaced sunflower plants growing under field conditions. Both N supply (range 0.25 to 11.25 g added N per plant) and leaf position significantly (p=0.001) affected maximum leaf area (LA_max) of target leaves through variations in leaf expansion rate (LER); effects on duration of expansion were small. Specific leaf nitrogen (SLN, g N m^-2) fell quite rapidly during the initial leaf expansion phase (LA < 35% LA_max) but leveled off during the final 65% increase of leaf area. This pattern held across leaf positions and N supply levels. Leaf nitrogen accumulation after 35% LA_max continued up to achievement of LA_max; reductions in the higher SLN characteristic of the initial phase were insufficient to cover the nitrogen requirements for expansion during the final phase. LER in the quasi-linear expansion phase (35 to 100% of LA_max) was strongly associated with SLN above a threshold that varied with leaf position (mean 1.79±0.225 g N m^-2). This contrasts with the response of photosynthesis at high irradiance to SLN, which has previously been shown to have a threshold of 0.3 g N m^-2; in the present work saturation of photosynthetic rate was evident when SLN reached 1.97 g N m^-2. Thus, once the area of a leaf exceeds 35% of LA_max, expansion proceeds provided SLN values are close to the levels required for maximum photosynthesis. However, growth of leaves during the initial expansion phase ensures a minimum production of leaf area even at low N supply levels.     

Xylem water flow in tropical vines as measured by a steady state heating method

Summary A method for determining the mass flow rate of xylem water in thin stems under natural field conditions is presented. Diurnal courses of xylem water flow and stomatal conductance of the vines Entadopsis polystachya, Cyclanthera multifoliolata, and Serjania brachycarpa were examined in a tropical deciduous forest on the west coast of Mexico. E. polystachya (leaf area 23.6 m²) had a maximum water flow rate of 6.50 kg h^-1 or 1.44 kg cm^-2 _{stem basal area} h^-1; daily water use was 2.00 kg m^-2 _{leaf area} day^-1. S. brachycarpa (leaf area 4.5 m²) and C. multifoliolata (leaf area 3.6 m²) had a maximum water flow rate of 0.72 and 0.19 kg h^-1 or 0.63 and 0.92 kg cm^-2 _{stem basal area} h^-1. Daily water use was 1.26 and 0.39 kg m^-2 _{leaf area} day^-1, respectively. The daily courses of xylem water flow were strongly influenced by the orientation of the leaf area to irradiance and its intensity. While leaves of E. polystachya had a constant high stomatal conductance during the day, S. brachycarpa had a maximum stomatal opening in the morning followed by continuous closure during the rest of the day. In contrast to the woody species, the herbaceous C. multifoliolata exhibited a strong midday depression of stomatal conductance and wilting of its leaves. The leaf biomass accounted for 8% (Entadopsis), 16% (Serjania), and 23% (Cyclanthera) of above-ground biomass. The relation of sapwood area to leaf area supplied (Huber value) was 0.19 (Entadopsis), 0.18 (Serjania), and 0.06 (Cyclanthera) cm² m^-2