Fertile forests produce biomass more efficiently.

Trees with sufficient nutrition are known to allocate carbon preferentially to aboveground plant parts. Our global study of 49 forests revealed an even more fundamental carbon allocation response to nutrient availability: forests with high-nutrient availability use 58 ± 3% (mean ± SE; 17 forests) of their photosynthates for plant biomass production (BP), while forests with low-nutrient availability only convert 42 ± 2% (mean ± SE; 19 forests) of annual photosynthates to biomass. This nutrient effect largely overshadows previously observed differences in carbon allocation patterns among climate zones, forest types and age classes. If forests with low-nutrient availability use 16 ± 4% less of their photosynthates for plant growth, what are these used for? Current knowledge suggests that lower BP per unit photosynthesis in forests with low- versus forests with high-nutrient availability reflects not merely an increase in plant respiration, but likely results from reduced carbon allocation to unaccounted components of net primary production, particularly root symbionts.

Europe-wide reduction in primary productivity caused by the heat and drought in 2003.

Future climate warming is expected to enhance plant growth in temperate ecosystems and to increase carbon sequestration. But although severe regional heatwaves may become more frequent in a changing climate, their impact on terrestrial carbon cycling is unclear. Here we report measurements of ecosystem carbon dioxide fluxes, remotely sensed radiation absorbed by plants, and country-level crop yields taken during the European heatwave in 2003. We use a terrestrial biosphere simulation model to assess continental-scale changes in primary productivity during 2003, and their consequences for the net carbon balance. We estimate a 30 per cent reduction in gross primary productivity over Europe, which resulted in a strong anomalous net source of carbon dioxide (0.5 Pg C yr(-1)) to the atmosphere and reversed the effect of four years of net ecosystem carbon sequestration. Our results suggest that productivity reduction in eastern and western Europe can be explained by rainfall deficit and extreme summer heat, respectively. We also find that ecosystem respiration decreased together with gross primary productivity, rather than accelerating with the temperature rise. Model results, corroborated by historical records of crop yields, suggest that such a reduction in Europe's primary productivity is unprecedented during the last century. An increase in future drought events could turn temperate ecosystems into carbon sources, contributing to positive carbon-climate feedbacks already anticipated in the tropics and at high latitudes.

The influence of climate and fructification on the inter-annual variability of stem growth and net primary productivity in an old-growth, mixed beech forest.

The periodic production of large seed crops by trees (masting) and its interaction with stem growth has long been the objective of tree physiology research. However, very little is known about the effects of masting on stem growth and total net primary productivity (NPP) at the stand scale. This study was conducted in an old-growth, mixed deciduous forest dominated by Fagus sylvatica (L.) and covers the period from 2003 to 2007, which comprised wet, dry and regular years as well as two masts of Fagus and one mast of the co-dominant tree species Fraxinus excelsior (L.) and Acer pseudoplatanus (L.). We combined analyses of weather conditions and stem growth at the tree level (inter- and intra-annual) with fruit, stem and leaf production, and estimates of total NPP at the stand level. Finally, we compared the annual demand of carbon for biomass production with net canopy assimilation (NCA), derived from eddy covariance flux measurements, chamber measurements and modelling. Annual stem growth of Fagus was most favoured by warm periods in spring and that of Fraxinus by high precipitation in June. For stem growth of Acer and for fruit production, no significant relationships with mean weather conditions were found. Intra-annual stem growth of all species was strongly reduced when the relative plant-available water in soil dropped below a threshold of about 60% between May and July. The inter-annual variations of NCA, total NPP and leaf NPP at the stand level were low (mean values 1313, 662 and 168 g C m(-2) year(-1), respectively), while wood and fruit production varied more and contrarily (wood: 169-241 g C m(-2) year(-1); fruits: 21-142 g C m(-2) year(-1)). In all years, an annual surplus of newly assimilated carbon was calculated (on average 100 g C m(-2) year(-1)). The results suggest that stem growth is generally not limited by insufficient carbon resources; only in mast years a short-term carbon shortage may occur in spring. In contrast to common assumption, stem growth alone is not a sufficient proxy for total biomass production or the control of carbon sequestration by weather extremes.

Air Pollution and Forest Decline in a Spruce (Picea abies) Forest.

Symptoms of forest decline of spruce in Europe range from needle yellowing and loss to tree and stand mortality. In a study area in northeast Bavaria, West Germany, where forest decline was initially detected, exposure to high concentrations of gaseous pollutants, SO(2,) NO(x,) and ozone has had no long-lasting direct effect on needles, and pathogens have only been secondary agents. Deposition of sulfur, nitrate, and ammonium, however, have significantly modified plant nutrition and soil chemistry. Spruce roots apparently take up ammonium rather than nitrate with an antagonistic effect on uptake of Mg. Nitrate left in the soil solution is leached together with sulfate to ground water, accelerating soil acidification and decreasing Ca/Al and Mg/Al ratios in the soil solution. Soil solution chemistry affects root development, and water and nutrient uptake. Had all nutrients become equally deficient, spruce trees probably could have adjusted by retarding their growth. However, canopy uptake of atmospheric nitrogen in addition to root uptake stimulated growth and caused a nitrogen to cation imbalance to develop; this imbalance resulted in the decline symptoms.

Xylem-tapping mistletoes: water or nutrient parasites?

Most mistletoes parasitize higher plants by tapping the xylem (a conduction tissue) of their hosts. Field observations of diurnal gas exchange parameters and carbon isotope ratios in xylem-tapping mistletoes from three continents support the hypotheses that water use efficiency and carbon isotope composition are related and that mistletoes which are parasitic for water are also nutrient parasites, differing in their water use efficiency relative to that of their hosts on the basis of host nitrogen supply in the transpiration stream.

Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient.

Several biodiversity experiments have shown positive effects of species richness on aboveground biomass production, but highly variable responses of individual species. The well-known fact that the competitive ability of plant species depends on size differences among species, raises the question of effects of community species richness on small-stature subordinate species. We used experimental grasslands differing in species richness (1-60 species) and functional group richness (one to four functional groups) to study biodiversity effects on biomass production and ecophysiological traits of five small-stature herbs (Bellis perennis, Plantago media, Glechoma hederacea, Ranunculus repens and Veronica chamaedrys). We found that ecophysiological adaptations, known as typical shade-tolerance strategies, played an important role with increasing species richness and in relation to a decrease in transmitted light. Specific leaf area and leaf area ratio increased, while area-based leaf nitrogen decreased with increasing community species richness. Community species richness did not affect daily leaf carbohydrate turnover of V. chamaedrys and P. media indicating that these species maintained efficiency of photosynthesis even in low-light environments. This suggests an important possible mechanism of complementarity in such grasslands, whereby smaller species contribute to a better overall efficiency of light use. Nevertheless, these species rarely contributed a large proportion to community biomass production or achieved higher yields in mixtures than expected from monocultures. It seems likely that the allocation to aboveground plant organs to optimise carbon assimilation limited the investment in belowground organs to acquire nutrients and thus hindered these species from increasing their performance in multi-species mixtures.

Cytokinins in the xylem sap of desert-grown almond (Prunus dulcis†) trees: Daily courses and their possible interactions with abscisic acid and leaf conductance.

Xylem sap samples were obtained from one- to four-year-old almond trees [Prunus dulcis (Miller) D. A. Webb] that had been grown in lysimeters of different volumes with different amounts of available water and subjected to an annual drying cycle. The samples were analyzed for cytokinin patterns by ELISA during the growing season. Free bases, ribosides and nucleotides of the N6 -(Δ2 -isopentenyl) adenine (iP) and the zeatin (Z) type could be identified and quantified. Z-type cytokinins were always predominant. In many cases the concentrations of the cytokinin fractions were rather constant during the day. In a limited number of days, however, Z-type cytokinins showed peak concentrations in the morning and a rapid decrease in the afternoon. A correlation between water status of the trees and the concentrations of cytokinins or their daily variations in the xylem sap could not be established. When the concentration of abscisic acid in the xylem sap was not limiting leaf conductance, daily-variation of a cytokinin fraction preceded daily variation in leaf conductance. We conclude that in almond trees, cytokinins may affect stomatal behaviour on a short-term basis. This is an outcome of changes in their xylem sap concentrations during the course of a day. Abscisic acid, on the other hand, acts as an opposing signal, the size of which reflects long-term water deficit.

A new type of climatized gas exchange chamber for net photosynthesis and transpiration measurements in the field.

A temperature- and humidity-controlled plant chamber for CO2 and H2O exchange measurements in the field is described in which the heat exchanger assembly and humidity controlling water vapour trap are separated from the plant cuvette. The shape and construction material of the plant cuvette can vary according to the demands of the experimental conditions and the size and growth form of the plant. The natural illumination field is only slightly altered in this plant cuvette. In the chamber, the temperature and humidity conditions can either be held constant throughout a wide range of conditions or can be programmed to track ambient condition. In this manner, not only temperature and absolute humidity are replicated, but it is also possible to reproduce the natural conditions of water vapour gradient between the evaporating surfaces in the mesophyll and the atmosphere, the relative humidity of the air, and the temperature difference between the leaf and the ambient air. Thus, the chamber appears to be an appropriate instrument to investigate with sufficient accuracy the reactions of individual plants in cultivation or in natural communities under field conditions.

[The effect of light and temperature of the CO2 exchange of different life forms in the ground vegetation of a montane beech forest]. / Die wirkung von licht und temperatur auf den CO2-gaswechsel verschiedener lebensformen aus der krautschicht eines montanen buchenwaldes.

In a montane beech (Fagus sylvatica) forest the influence of the climatic factors, light and temperature, on net photosynthesis and on the CO2 balance of the ground vegetation was investigated. The total turnover of carbon was calculated. Species studied included: Athyrium filix-femina, Oxalis acetosella, Luzula luzuloides, Deschampsia flexuosa and young plants of Fagus sylvatica. 1. The light compensation point in all spp. is between 300 and 500 lux except for D. flexuosa where it is 2 klx. Light saturation is attained at 2-3 klx for A. filix-femina, at 5-6 klx for O. acetosella, and at 6-7 klx for L. luzuloides and F. sylvatica. The net photosynthesis of D. flexuosa increases linearly upto 12 klx. This plant, therefore, is more closely related to plants with high light requirements than all the other species under experiment. 2. The maximum rates of net photosynthesis in O. acetosella and A. filix-femina are higher than in all the other plants, independent of the reference system. Per unit dry weight they even attain rates of CO2 uptake (22-27 mg CO2/gdw·h) known from herbs under the much better light conditions of an open habitat. F. sylvatica and L. luzuloides exhibit per unit dry weight only 30% of this rate and D. flexuosa 25%. On a leaf surface area and chlorophyll content basis differences are smaller: F. sylvatics attains 75%, L. luzuloides reaches 50% and D. flexuosa only 30% of the maximal rates of net photosynthesis of O. acetosella and A. filix-femina. The higher CO2 uptake of O. acetosella and A. filix-femina points to a better adaptation of their photosynthetic apparatus in comparison to all the other species of the same habitat. 3. At light saturation the temperature optimum of A. filix-femina and O. acetosella covers a smaller range at lower temperatures than was found in the other species. These attain almost maximal rates of net photosynthesis over the whole range of temperatures of their natural habitat. At decreasing light intensities the temperature optimum of O. acetosella changes from 13-18° C at 8-12 klx to a range of even lower temperatures (9-12° C at 1 klx). 4. The respiration of the rhizome and the roots of O. acetosella is per unit dry weight 40% of the dark respiration rate in the above ground material. 5. The daily gain of net photosynthesis per unit dry weight of O. acetosella and A. filix-femina is 4 times as high as in L. luzuloides and in F. sylvatica and 7 times as high as in D. flexuosa. Per unit of surface area and chlorophyll content differences are smaller. The sequence in all cases remains the same. During the night D. flexuosa has the highest relative respiratory loss. Its CO2 gain over 24 hours is very small. 6. The importance of sun flecks on the CO2 balance is small in all species except D. flexuosa. More important is the mean light intensity and the rate of net photosynthesis which is attained under these conditions. The amount of CO2 photosynthetically bound in sun flecks is 6% of the daily balance in A. filix-femina, 16-19% of the daily balance in O. acetosella, L. luzuloides and F. sylvatica, and 27% of the daily balance in D. flexuosa. The existence of D. flexuosa is dependent on the occurrence of sun flecks on the forest floor. The ecological significance of the relative light intensity in the mosaic-like distribution of plants on the forest floor is discussed. 7. The varying success in adaptation to the conditions of the habitat becomes even more evident when compared with the primary production of the beech crown. The daily gain of net photosynthesis of O. acetosella and A. filix-femina per unit dry weight is much larger than in either the sun or shade leaves in the canopy of the same stand. Per unit surface area of the leaves they attain 18-20%, per unit chlorophyll content 32% (L. luzuloides and the young plants of F. sylvatica 16-27%, D. flexuosa 4%) of the gain of net photosynthesis in the beech sun leaves. 8. A comparison with a model of primary production (maximal rates of net photosynthesis under experimentally optimal conditions over the whole day = 100%) shows what effect the different climatic factors of the natural habitat have in limiting the CO2 balance, and to what extend the actual CO2 gain reaches the physiological optinum. On the forest floor the rate of net photosynthesis is reduced primarily through the intense shade of the beech canopy and by dawn and dusk (reduction of the maximal CO2 gain in O. acetosella and in A. filix-femina ca. 50%, in L. luzuloides and F. sylvatica ca. 60% and in D. flexuosa ca. 86%). The effect of additional clouds is smaller (reduction of the maximal CO2 gain in all species 4-6%, maximal 19%). The effect of temperature is very small for L. luzuloides, F. sylvatica and D. flexuosa. Corresponding to the low temperature optimum the influence of the prevailing temperatures is much higher in O. acetosella and A. filix-femina (4-6% reduction of the maximal CO2 gain through temperatures above optimum). 9. For an assessment of the competition potential of O. acetosella compared to L. luzuloides and F. sylvatica for the vegetation period of April to August, the carbon balances of the whole plants were estimated. These are compared with the dry weight increase. Among the three life forms of the rhizome geophyte (Oxalis), the hemicryptophyte (Luzula) and the phanerophyte (Fagus) there are striking differences in the use and in the distribution of the CO2 gain. L. luzuloides invests 65% of the net photosynthetic gain as dry weight increment (O. acetosella only 44%, F. sylvatica 40%). Moreover, the growth of L. luzuloides takes place primarily above ground with the establishment of new leaves. The relative proportion of the growth above ground to the total dry weight increment is for L. luzuloides 63%, as against 57% in F. sylvatica and only 42% in O. acetosella. In respect to the total carbon balance the better use of the CO2 gain gives L. luzuloides in this habitat a higher competition potential than the photosynthetically more active O. acetosella. The constitutional differences in the photosynthetic activity are compensated for by the distribution of the assimilates to shoot and root.

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

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-2stem basal area h-1; daily water use was 2.00 kg m-2leaf 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-2stem basal area h-1. Daily water use was 1.26 and 0.39 kg m-2leaf 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.

Insect capture and growth of the insectivorous Drosera rotundifolia L.

Rates of insect capture increased with leaf area in the insectivorous plant Drosera rotundifolia, and growth of new leaves was related to insect capture. However, increased leaf growth was counterbalanced by leaf abscission which was in turn related to insect capture and leaf growth. Leaf loss equaled leaf growth in plants having natural rate of insect capture. A large proportion of the nitrogen gain from prey was stored in the hypocotyl; it was estimated from feeding experiments that about 24% to 30% of the nitrogen stored in the hypocotyl after winter originated from insect capture in the previous season. The effect of insect capture is discussed in relation to the life cycle of Drosera.

The effect of nitrogen nutrition on growth and biomass partitioning of annual plants originating from habitats of different nitrogen availability.

The hypothesis was tested that faster growth of nitrophilic plants at high nitrogen (N) nutrition is counterbalanced by faster growth of non-nitrophilic plants at low N-nutrition. Ten annual plant species were used which originated from habitats of different N-availability. The species' preference for N was quantified by the "N-number" of Ellenberg (1979), a relative measure of nitrophily. The plants were cultivated in a growth cabinet at five levels of ammonium-nitrate supply. At low N-supply, the relative growth rate (RGR) was independent of nitrophily. At high N-supply, RGR tended to be higher in nitrophilic than in non-nitrophilic species. However, the response of RGR to N-supply was strongly and positively correlated with the nitrophily of species. Increasing N-supply enhanced partitioning to leaf weight per total biomass (LWR) and increased plant leaf area per total biomass (LAR). Specific leaf weight (SLW) and LWR were both higher in non-nitrophilic than in nitrophilic species at all levels of N-nutrition. NAR (growth per leaf area or net assimilation rate) increased with nitrophily only under conditions of high N-supply. RGR correlated positively with LAR, irrespective of N-nutrition. Under conditions of high N-supply RGR correlated with SLW negatively and with NAR positively.

Carbon and nitrogen isotope ratios in different compartments of a healthy and a declining Picea abies forest in the Fichtelgebirge, NE Bavaria.

Natural carbon and nitrogen isotope ratios were measured in different compartments (needles and twigs of different ages and crown positions, litter, understorey vegetation, roots and soils of different horizons) on 5 plots of a healthy and on 8 plots of a declining Norway spruce (Picea abies (L.) Karst.) forest in the Fichtelgebirge (NE Bavaria, Germany), which has recently been described in detail (Oren et al. 1988a; Schulze et al. 1989). The δ13C values of needles did not differ between sites or change consistently with needle age, but did decrease from the sun-to the shade-crown. This result confirms earlier conclusions from gas exchange measurements that gaseous air pollutants did no long-lasting damage in an area where such damage was expected. Twigs (δ13C between-25.3 and-27.8‰) were significantly less depleted in 13C than needles (δ13C between-27.3 and-29.1‰), and δ13C in twigs increased consistently with age. The δ15N values of needles ranged between-2.5 and-4.1‰ and varied according to stand and age. In young needles δ15N decreased with needle age, but remained constant or increased in needles that were 2 or 3 years old. Needles from the healthy site were more depleted in 15N than those from the declining site. The difference between sites was greater in old needles than in young ones. This differentiation presumably reflects an earlier onset of nitrogen reallocation in needles of the declining stand. δ15N values in twigs were more negative than in needles (-3.5 to-5.2‰) and showed age- and stand-dependent trends that were similar to the needles. δ15N values of roots and soil samples increased at both stands with soil depth from-3.5 in the organic layer to +4‰ in the mineral soil. The δ15N values of roots from the mineral soil were different from those of twigs and needles. Roots from the shallower organic layer had values similar to twigs and needles. Thus, the bulk of the assimilated nitrogen was presumably taken up by the roots from the organic layer. The problem of separation of ammonium or nitrate use by roots from different soil horizons is discussed.

Environmental control of crassulacean acid metabolism in Welwitschia mirabilis Hook. Fil. in its range of natural distribution in the Namib desert.

Within the area of its natural distribution in South West Africa, Welwitschia mirabilis has a less negative δ13C value than C3 plants and a more negative δ13C value than C4 species. This indicates that Welwitschia m. assimilates CO2 partially via CAM when growing in its natural habitat. The difference between the δ13C values of Welwitschia m. and of the C3 species is significant in the savanna, whereas it is only small and statistically not significant in the grassland zone. The proportion of CO2 fixed via CAM is largest in the coastal desert zone. There was no correlation between the δ13C values and the Cl- or ash content of the tissue. Thus, CAM in Welwitschia m. seems not to be induced by salt stress. There is no change in the δ13C values along the persistent Welwitschia m. leaf. The present data indicate that on a broad geographical scale in the area of distribution temperature regime, and water stress as a modifying factor, determine CAM in Welwitschia m. The ecological implications are discussed by comparing the behaviour of Welwitschia m. with other CAM, C3 and C4 species of the accompanying flora.

Spacial distribution of photosynthetic capacity and performance in a mountain spruce forest of Northern Germany : I. Biomass distribution and daily CO2 uptake in different crown layers.

Biomass distribution and diurnal CO2 uptake under natural conditions were investigated on Picea abies in a mountainous climate (Solling, Northwest Germany). Spruce has a remarkable variability in leaf characteristics. Even on a single branch in the lower sun crown, needle dry weight and surface area change considerably from the branch base to the tip and accoring to exposure. Only about 18% of the total biomass of the tree was current year's growth, about 40% of the needles were 4 years and older reaching a maximal age of 12 years. The main growing zone was at the border of upper shade and lower sun crown and the main accumulation of dry weight was at a greater tree height than was observed for maximal growth of needle numbers or surface area. The annual, new growth shifted toward the upper sun crown. Maximal daily CO2 uptake was highest in the lower sun crown on days with variable cloud cover when temperatures were moderate and water vapor pressure deficits were low. Also the annual CO2 uptake was highest in the lower sun crown, where 4-year-old and older needles contributed about 35% to the annual CO2 uptake of the tree. Current year growth contributed about 15% of the total CO2 gain. The upper and lower sun crowns produce about 70% of the total carbon gain. The carbon balance of spruce and the distribution of the production process in relation to needle age and crown level are discussed.

Spacial distribution of photosynthetic capacity and performance in a mountain spruce forest of northern Germany : II. Climatic Control of Carbon Dioxide Uptake.

Net photosynthesis of Picea abies was measured in a spruce forest in northern Germany with temperature- and humidity-controlled cuvettes in 4 different crown layers on shoots of different ages. These measurments were performed such that temperature and humidity either followed ambient conditions or were kept constant. Annual courses of light-, temperature-, and humidity-related net photosynthesis were determined. Spruce had a remarkably constant rate of CO2 uptake from April to September for 1-year and older needles. Light saturation was achieved at 25 klx. Current year needles had the highest rates of CO2 uptake in early summer, but these rates decreased by autumn. Photosynthetic capacity decreased with needle age and, on a dry weight basis, it was higher in the shade than in the sun crown. The temperature optimum was between 13 and 23° C. Photosynthesis in spruce decreased when air humidity was low.The effect of the natural weather conditions on photosynthetic capacity was determined. The habitat is characterized by a high frequency of low light intensities (75% of total daytime below 20 klx) and cool temperatures (80% of daytime between 9 and 21° C). Low air humidity was only present when light intensities were high. The major limiting factor for production was low light intensities, which reduced photosynthetic capacity in the sun crown to 42% below maximum possible rates. Adverse temperatures reduced CO2 uptake by 28% and large water vapor pressure deficits reduced rates by only 2% compared with maximum possible rates. The limited adaptation to light is discussed.

Spacial distribution of photosynthetic capacity and performance in a mountain spruce forest of Northern Germany : III. The significance of the evergreen habit.

Growth and CO2 uptake in the crown of a spruce tree is described and the production processes of this evergreen conifer are compared with those of a deciduous beech. Spruce had 60% lower rates of net photosynthesis per dry weight than beech. But, beech had a 30% shorter growing season and a 84% smaller biomass than spruce. The annual CO2 gain was 40% lower in beech than it was in spruce.An analysis shows the following conclusions for this habitat. (1) The effect of a prolonged growing season is small. The annual CO2 gain of spruce would be reduced only by 9% if the growing season was the same length as for beech. (2) The annual CO2 gain would increase 14% if all needles in spruce were deciduous, because the current year needles have a higher average rate of CO2 uptake than 3-year old and older needles, but a lower average rate than 1- and 2-year old ones. However, the carbon balance of the tree shows that spruce could not afford to produce the existing needle biomass (14 t ha-1) each year. (3) If spruce were to produce the same deciduous foliage biomass during the same growing season as beech then total production by spruce would be reduced 67%. (4) The annual CO2 uptake by evergreen spruce was higher than deciduous beech not because of a long growing season, but because of the longevity of its needles, which during their total life time (an average of 5 years) have a two to three times greater CO2 uptake than a deciduous leaf in one summer season. The relatively small investment in current year needles produces an annually low, but long lasting assimilation of CO2.

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content : II. In the mesophytic herbaceous species Helianthus annuus.

The responses of leaf water potential, leaf conductance, transpiration rate and net photosynthetic rate to vapour pressure deficits varying from 10 to 30 Pa kPa-1 were followed in Helianthus annuus as the extractable soil water decreased. With a vapour pressure deficit of 25 Pa kPa-1 around the entire plant as the soil water content decreased, the leaf conductance and transpiration rate showed a strong closing response to leaf water potential at a value of-0.65 MPa, whereas with a vapour pressure deficit of 10 Pa kPa-1 around the entire plant, the rate of transpiration and leaf conductance decreased almost linearly as the leaf water potential decreased from-0.4 to-1.0 MPa. Increasing the vapour pressure deficit from 10 to 30 Pa kPa-1 in 5 Pa kPa-1 steps decreased the leaf conductance by a similar proportion at all extractable soil water contents. At high soil water contents, the decrease in conductance with leaf water potential was greater when the vapour pressure deficit was increased than when it was not, indicating a direct influence of vapour pressure deficit on the stomata. The rate of net photosynthesis decreased to a smaller degree than the leaf conductance when the vapour pressure deficit around the leaf was varied. Overall, the net photosynthetic rate decreased almost linearly from 20 to 25 µmol m-2 s-1 at-0.3 MPa to 5 µmol m-2 s-1 at-1.2 MPa. As the soil water decreased, the internal carbon dioxide partial pressure was maintained between 14 and 25 Pa.No unique relationship between leaf conductance, transpiration rate or photosynthetic rate and leaf water potential was observed, but in all experiments leaf conductance and the rate of net photosynthesis decreased when about two-thirds of the extractable water in the solid had been utilized irrespective of the leaf water potential. We conclude that soil water status, not leaf water status, affects the stomatal behaviour and photosynthesis of H. annuus.

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content : III. In the sclerophyllous woody species Nerium oleander.

The responses of leaf conductance, leaf water potential and rates of transpiration and net photosynthesis at different vapour pressure deficits ranging from 10 to 30 Pa kPa-1 were followed in the sclerophyllous woody shrub Nerium oleander L. as the extractable soil water content decreased. When the vapour pressure deficit around a plant was kept constant at 25 Pa kPa-1 as the soil water content decreased, the leaf conductance and transpiration rate showed a marked closing response to leaf water potential at-1.1 to-1.2 MPa, whereas when the vapour pressure deficit around the plant was kept constant at 10 Pa kPa-1, leaf conductance decreased almost linearly from-0.4 to-1.1 MPa. Increasing the vapour pressure deficit from 10 to 30 Pa kPa-1 in 5 Pa kPa-1 steps, decreased leaf conductance at all exchangeable soil water contents. Changing the leaf water potential in a single leaf by exposing the remainder of the plant to a high rate of transpiration decreased the water potential of that leaf, but did not influence leaf conductance when the soil water content was high. As the soil water content was decreased, leaf conductances and photosynthetic rates were higher at equal levels of water potential when the decrease in potential was caused by short-term increases in transpiration than when the potential was decreased by soil drying.As the soil dried and the stomata closed, the rate of photosynthesis decreased with a decrease in the internal carbon dioxide partial pressure, but neither the net photosynthetic rate nor the internal CO2 partial pressure were affected by low water potentials resulting from short-term increases in the rate of transpiration. Leaf conductance, transpiration rate and net photosynthetic rate showed no unique relationship to leaf water potential, but in all experiments the leaf gas exchange decreased when about one half of the extractable soil water had been utilized. We conclude that soil water status rather than leaf water status controls leaf gas exchange in N. oleander.