Current issue: 58(4)
How will global warming affect southern populations of boreal trees? In paper birch, Betula papyrifera (Betulaceae), alpine trees with an evolutionary history of relatively cool summers may be more sensitive to climate warming than valley populations. We evaluated this scenario by growing seedlings from different populations in four temperature treatments (mountain field site, valley field site, and two greenhouse rooms).
Populations from low elevations germinated earlier and had higher germination success than population from high elevations (16.8 vs. 22.0 d; 72% vs. 11%). At the valley site, seedlings from native populations grew faster than seedlings from higher elevations (mean ± SE = 0.25 ± 0.02 vs. 0.09 ± 0.04 mm · cm-1 · d-1) while at the mountain site, all seedlings grew at similar rates. Seedling grown in cooler environments had higher root : shoot ratios, perhaps to compensate for temperature limitations in nutrient uptake by roots. Leaf area varied among populations but was not affected by environmental differences across the field sites. Net photosynthetic rates at valley temperatures were higher for seedlings grown in the valley than for seedling grown in the mountains or the warm greenhouse (12.0 vs. 10.3 and 5.8 μmoles · m-2 · s-1), perhaps due to adaptive phenotypic adjustments. Climatic warming could rapidly produce important phenotypic changes in birch trees (e.g. decreased root : shoot ratio, reduced growth in alpine populations). On a longer time-scale, warming could also result in genetic changes as natural selection favours valley genotypes in alpine sites where they are presently rare.
A total of 1,800 3-year old seedlings of Norway spruce (Picea abies (L.) H. Karst.) from two Norwegian and one German provenance were treated with two different nitrogen levels during the 1992 growth season. The plants were kept during the following winter at two different temperature levels. In the spring of 1993, the nutrient application was resumed, and the plants were divided between three different treatments, 350 and 650 p.p.m. in open top chamber and a control plot outside the chambers. This treatment was repeated also during the following 1994 growth season.
The growth and primary production were studied by photosynthesis experiments and by non-destructive growth measurements. The result indicate that raised winter temperature may lead to increased needle loss and reduced growth the following season, particularly in northern provenances. Carbon dioxide significantly influenced growth in addition to nutrient level and winter temperature. High CO2 also seemed to cause increased photosynthesis at early season, and earlier budbreak and growth cessation than in control plants.
Change in dry matter partitioning, 14C-incorporation, and sink 14C-activity of 1.5-year-old Scots pine (Pinus sylvestris L.) seedlings grown in growth chamber conditions were studied during a 91-day experiment. On five sampling dates, seedlings were labelled with 14CO2 and whole-plant allocation patterns were determined. Intensively growing shoots modified the dry matter partitioning: during shoot growth the proportion of roots decreased but after that it increased. Based on their large proportion of dry matter, the needles (excluding current needles) were the strongest sink of carbon containing 40% of the incorporated 14C. Despite their small initial sink size, the elongating shoots (current main shoot + current branch) and their needles were the second strongest sink (30–40% of the total 14C) which reflects their high physiological activity. The proportion of 14C in the current year’s main shoot increased during shoot growth but decreased as the growth began to decline after 70 days. 10–20% of the total assimilated 14C was translocated to the roots. Laterals above 2nd order were the strongest sink in the root system, containing twice as much 14C as the other roots together. Alternation between shoot and root growth can be seen clearly: carbon allocation to roots was relatively high before and after the period of intensive shoot growth. Changes in root sink strength resulted primarily from changes in root sink activity rather than sink size.
The Värriö environmental measurement station has been designed and constructed during 1991 and 1992. The measurement system consists of measurement units for gases (sulphur dioxide, ozone, carbon dioxide), particles, photosynthesis and irradiation. A meteorological station is also included. The preliminary measurement period was started on August, 1991. During the first year (1991–1992) some parts of the system were redeveloped and rebuilt. Full, continuous measurement started in August 1992. The system has been working quite reliably, with good accuracy. The preliminary results show that pollution episodes are observed when the wind direction is from Monchegorsk or Nikel, the main emission sources in Kola Peninsula.
The interactive effects of potassium deficit and foliar application with acid water (pH 5.5, 4.5, 4.0 and 3.0 given consecutively) on CO2 exchange rate of Pinus sylvestris L. seedlings was investigated in field conditions. No reduction of the CO2 exchange rate was observed in the seedlings supplied with sufficient potassium. Only the seedlings having the lowest needle K concentration (2.4 mgg-1) had an apparently low CO2 exchange rate before the applications with acid water. The CO2 exchange rate of most of the seedlings with low needle K concentration (3.9–6.0 mgg-1) decreased after the acid water application. The threshold acidity for the reduction varied between pH 4,0 and 3.0 depending on the needle K concentration. The reduction was more apparent at high irradiance. It was concluded that acid precipitation disturbs the CO2 exchange only in conditions of mineral nutrient deficit.
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A close relationship between photosynthetic capacity and nitrogen concentration of leaves is known to exist. In conifers, nitrogen also affects the pattern of mutual shading within a shoot, which is a basic unit used in studying photosynthesis of coniferous trees. These effects of needle nitrogen concentration on photosynthetic capacity and mutual shading of needles were analysed for Scots pine (Pinus sylvestris L.) shoots taken from five young stands growing on sites of different fertility. The effect of nitrogen concentration on needle photosynthesis was studied based on measurements of the photosynthetic radiation response of shoots from which two thirds of the needles were removed in order to eliminate the effect of within shading.
An increase of one percentage unit in nitrogen concentration of needles increased the photosynthetic capacity of needles by 25 mg CO2 dm-2h-1. The effect of nitrogen on within-shoot shading was quantified in terms of the silhouette area to total needle area ratio of a shoot (STAR), which determines the relative interception rate per unit of needle area on the shoot. Although nitrogen promoted needle growth, an increase in nitrogen concentration decreased the within-shoot shading. This effect resulted from a decrease in needle density on the shoot and an increased needle angle with increasing nitrogen content.
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A third generation of forest tree gas exchange measuring system design for the use in the field is described. The system is designed to produce data for determining the dependence of the rate of tree photosynthesis, respiration and transpiration on environmental factors. The system consists of eight cuvettes, a tubing system, two infrared gas analysers, an air flow controller, a data logger, and a computer. The measuring cuvette is a clap type, i.e. it is mostly open, only closing during measurement. CO2 exchange is measured as the change in the cuvette concentration of CO2, and, transpiration is measured as the increase in water vapour concentration while the cuvette is closed. The environmental factors measured are temperature, irradiance and air pressure. The system was planned in 1987 and constructed in 1988. It worked reliably in late summer 1988 and the quality of data seems to be satisfactory.
The PDF includes an abstract in Finnish.
A comparison study concerning the effects of acid rain on Scots pine (Pinus sylvestris L.) seedlings has been performed. Two different X-ray fluorescence methods, PIXE and IXRF, were employed to produce multielement analyses of the samples. Seedlings were treated for 3 months with watering of pH=7 or pH=3 liquids on the needles and on the roots. One year and two years old needles of the seedlings were inspected for changes in photosynthetic rate as well as for changes in elemental concentrations.
Twelve elements from Si to Zn were compared in the samples. The PIXE results show that the amounts of most of these elements in the needles of the seedlings grown in sand increase, when treated with acid water. This growth is clearer when the roots are treated with acid water. The elemental concentrations of the needles in the seedlings grown in soil on the other hand decrease slightly.
A model was constructed, the aim of which was to predict growth under conditions where air pollutants are present. The model is based on photosynthesis and on the allocation of photosynthetic products for growth. It is assumed that air pollutants released during energy production mainly affect photosynthesis in two ways: 1) directly by injuring the photosynthetic mechanism, and 2) indirectly by leaching nutrients. The two ways were studied empirically in order to identify a sub-model for the photosynthesis of a plant exposed to air pollutants.
The stand model will be applied to two purposes. The present stage of forests in Finland is compared with the simulated state based on the assumption that no pollutants are present. In addition, the decrease in forest yield under different conditions derived from predictions about long-range pollutant transport in Europe is analysed.
The crown structure and stem growth of Norway spruce (Picea abies (L.) H. Karst.) undergrowth was studied in relation to the prevailing light conditions and potential photosynthesis. Shading decreased the stem height growth more than the length increment of laterals, producing a plate-shaped crown in deep shade. Needles responded to shading by adopting a horizontal inclination in deep shade. The needles were wide and thin respectively in shade. In the open the needle cross-section was almost square. Stem radial growth and height growth were both affected by shading exhibiting a linear response to the prevailing light conditions and the potential photosynthesis. Light conditions under dominating trees were closely correlated with the basal area of the dominating trees.
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Needle damages, transpiration, photosynthesis and needle and stem height growth of Scots pine (Pinus sylvestris L.) seedlings treated with dilute sulphuric acid were studied. The acidity of the solution was pH 3. Application of a dilute solution of sulphuric acid equivalent to the normal amount of precipitation occurring during the growing season damaged the surface of two-year-old needles but not that of the current-year needles. A reduction in the photosynthetic rate of 10–30% was observed compared with the untreated seedlings. Transpiration of the seedlings was not affected by the treatment. Needle growth and stem height growth of the seedlings growing on a substrate representing poor sandy soil were reduced. Increased needle growth and stem height growth were characteristic for the seedlings growing on substrate representing fertile moraine.
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The technique of double normalizing, i.e. normalizing the relative needle biomass and the length of the living crown system, is applied to the modelling of the distribution of needle biomass in the canopy of young Scots pine (Pinus sylvestris L.) stands. The study based on the parameters of β-function shows that at the individual-tree level, the variance in needle distribution was not closely associated with any tree characteristics. A shift in the point of maximum needle biomass upwards unsuppressed trees was, however, evident. This was associated with an increase in the height of the trees. At the stand level, the stand mean height and stand density had an equal and a rather high potential for explaining the variance in the needle distribution. The normalized crowns are utilized in models for determining light extinction in the crown. A special technique for determining the amount of photosynthates available for growth in a particular tree is presented.
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The effect of solar radiation and air temperature on the basic density of Scots pine (Pinus sylvestris L.) wood has been investigated on the basis of material obtained from the literature. Solar radiation seemed to affect basic density during earlywood formation. Temperature had the greatest effect on basic density in late summer. The varying effects of radiation and temperature seemed to be associated with the dynamics of the crown system of trees. Especially the capacity of the crown system to produce the amount of photosynthesis needed in tracheid growth is assumed to be of importance in controlling the variation in the basic density of Scots pine wood. Growth of thracheids from the point of view of photosynthate supply is discussed.
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A system for measuring the net photosynthesis, transpiration and environmental factors within the canopy and ground cover vegetation is described. The system operates continuously throughout the growing season in a young Scots pine (Pinus sylvestris L.) stand. A data-logging unit controls the system and carries out the measurements on the readings of the sensors of photosynthesis, transpiration, light intensity outside the canopy, light climate inside the assimilation chambers, and dry and wet temperatures from selected points. These measurements are shown digitally and automatically punched onto paper tape.
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A quantitative method for determining the annual growth level of plant species has been presented. In particular, attention was paid to the dependence of the growth level on the amount of light available for photosynthesis. A mathematical model for the dependence of structural matter production on photosynthetic production has been presented for some plant species.
The study is based on the assumption that the total amount of annual net photosynthesis plays a role of primary importance in determining the relationship between photosynthetic production and structural matter production. The basic environmental factors determining the photosynthetic rate are light and temperature, if the water and nutrient supply is adequate. The dependence of photosynthetic rate on light and temperature was determined by monitoring the CO2 uptake rate of natural plant populations between the photosynthetic levels of different plant populations with an infrared gas analyser.
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The aim of the present paper was to study the annual production of Pleurozium schreberi (Brid.) Mitt., Hylocomnium splendens (Hedw.) B.S.G and Dicranum polysetum Sw. as a function of light available for photosynthesis. The productivity of the above moss species is studied using the harvested quadrats method in Norway spruce (Picea abies) stands of the Myrtillus site type representing different stand density classes (basal area from 0 to 34 m2/ha) in Southern Finland.
The annual production of each species in different stands was correlated with the amount of light available for photosynthesis i.e. with the photosynthetic production. Functions for the dependence of productivity on light conditions were produced for each species. The individual functions and their ecological significance is discussed. The adaptation of each species to low light intensity is evident since no meaningful addition to production takes place when the photosynthetic light ratio reaches values greater than 0.3–0.4. In other words, the level of photosynthesis which is 30–40% of that possible in the open, provides sufficient supply of carbohydrates or the basic functions of the moss species studied. Pleurozium schreberi and Dicranum polysetum seem to have greater light requirements than Hylocomnium splendens.
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The photosynthetic rate of Pleurosium schreberi (Willd.), Hylocomnium splendens (Hedw.) and Dicranum undulatum (Sw.) grown in plastic containers was monitored with infrared gas analyser in open air under natural weather conditions. It proved that the photosynthetic rate of wet moss cushions was satisfactorily predicted by temperature and light intensity. In dry moss cushions this kind of model gave too high an estimate for photosynthetic rate. Water requirements of each moss species were found to be moderate, and water content of moss cushions limited photosynthetic rate only under serious water deficiency.
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The activation of CO2 exchange was monitored in two Scots pine (Pinus sylvestris L.) seedlings transferred from the field to the laboratory in December. Gas exchange was monitored by an URAS I infrared gas analyser in a so-called open IRGA-system with trap type chambers. Transpiration was also measured at the same time by weighing the potted seedlings twice a day. The measuring period lasted eleven days. During the period, the level of both transpiration and net photosynthesis increased about ten times. Furthermore, it was found that the level of photosynthesis at high temperatures was relatively lower at the beginning than at the end of the measuring period.
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Light intensity inside the canopy varies considerably both in space and time. A new apparatus was developed which is disturbed as little as possible by the above-mentioned variation. The construction is based on the linear relationships between light intensity (measured using silicon diodes) and photosynthesis. This procedure permits linear operations (summing and integration) to be carried out on the output of the diodes without any loss of accuracy. There are five diodes in each assimilation chamber. A model, in which the independent variables include ligth, measured with the present equipment, and temperature, fits the photosynthetic rates well even inside the canopy.
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Photosynthesis and dark respiration in five families of autochtonous Norway spruce (Picea abies (L.) H. Karst.) and in seedlings from twenty Finnish stands of Scots pine (Pinus sylvestris L.) were investigated in constant environmental conditions. Values of CO2 exchange were compared with the height growth and weight of seedlings in Norway spruce and with the weight alone in Scots pine. No statistically significant differences were found in CO2 exchange among progenies or stands. Photosynthetic efficiency and photosynthetic capacity showed a positive correlation both in spruce and in pine. Growth and net photosynthetic capacity were linearly and positively correlated in pine. Spruce and a higher light compensation point than pine. The use of an open IRGA system with several simultaneous measurements and the trap-type cuvette construction in genetic work are discussed.
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In the literature review the current status of information on the genetic variation of CO2 exchange and some reviews and investigations on this subject are listed. Photorespiration is separately discussed and unpublished data of an electron microscope study of poplar leaf microbodies are presented.
Considerable genetic inter- and intraspecific variation is found in several characteristics that affect CO2 exchange in trees. Photosynthesis in young trees does not correlate well with growth through the whole rotation cycle. A special interest has been shown to marginal environmental conditions (e.g. water deficit, low temperature, and low light intensity), as opposed to optimal conditions often employed in laboratory studies of CO2 exchange in trees.
In an unpublished poplar studies by the author et.al. a preliminary experiment with poplar clones showed variation in the CO2 competition point. This variation was negatively correlated with the photosynthesis efficiency of these clones.
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Eight Scots pine (Pinus sylvestris L.) seed sources, ranging from 42° to 66° north latitude, were grown under a constant, 16-hour photoperiod in a greenhouse for approximately 6 months. Rates of photosynthesis, as measured by an IRGA, and growth, as measured by increase in height and fresh and dry weight, differed among seed sources at the end of the six-month growing period. Photosynthetic capacity and growth were strongly related to latitude of seed source, and were greatest in the seed sources coming from a parent environment in which maximum photoperiods are about 16 hours.
Photosynthetic efficiency (rate of photosynthesis per gram needle weight) was also strongly related to latitude of seed source, but was lowest in the seedlings which exhibited the greatest growth and photosynthetic capacity. This may have been due to (1) more mutual shading of needles on the larger seedlings and (2) a lesser proportion of juvenile needles on the larger seedlings or (3) biochemical differences in the use of photosynthate in the needles. Seed source and light intensity had an interacting effect on rates of photosynthesis only in seedlings of the two northernmost seed sources.
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The accompanying collective research report is the result of the research project in 1986–90 between the Finnish Academy and the former Soviet Academy of Sciences. The project was organized around common field work in Finland and in the former Soviet Union, and theoretical analyses of tree growth determining processes. Based on theoretical analyses, dynamic stand growth models were made and their parameters were determined utilizing the field results.
Annual cycle effects the tree growth. Our theoretical approach was based on adaptation to local climate conditions from Lapland to South Russia. The initiation of growth was described as a simple low and high temperature accumulation driven model. The model was linked with long-term temperature data.
Analysis of field measurements of CO2 exchange showed that irradiance is the dominating factor causing variation in photosynthetic rate in natural conditions during summer. The penetration of irradiance into Scots pine (Pinus sylvestris L.) canopy is a complicated phenomenon. A moderately simple but balanced forest radiation regime sub-model was constructed.
The common field excursions in different geographical regions resulted in a lot of experimental data of regularities of woody structure. The water transport seems to be a good common factor to analyse these properties of tree structure. The produced theoretical and experimental material was utilized in the development of stand growth model that describes the growth and development of Scots pine stands in Finland and the former Soviet Union. The core of the model is carbon and nutrient balances. This means that carbon obtained in photosynthesis is consumed or growth and maintenance and nutrients are taken according to the metabolic need. Since the fundamental metabolic processes are the same in all locations, the same growth model structure can be applied in the large range of Scots pine. The model enables the analysis of geographical differences in the growth of Scots pine. The present approach enables utilization of structural and functional knowledge gained in places of intensive research, in the analysis of growth and development of any stand.
The aim of this study was to investigate the ecophysiological and morphological characteristics of two salt-tolerant tree species, Eucalyptus camaldulensis Dehn. and Combretum quadrangulare Kurz. A greenhouse experiment with different levels of NaCl salinity (0, 0.5, 1.0, 1.5, and 2.0%) was set up and the results were compared with those of a field study on non-saline and saline soils. The determination of optimum gas exchange and the development and evaluation of photosynthetic models with and without water deficit were also included in this study.
Morphological characteristics under saline conditions showed that shoot height and diameter growth, shoot internode length, root length/biomass, leaf width and length, leaf area, number and biomass, and shoot/root and leaf/root ratios decreased with salinity, while leaf thickness increased with salinity. More growth was allocated to the roots than to the leaf canopy. Ecophysiological studies in laboratory showed that photosynthesis, stomatal conductance and water potential decreased with salinity, while the CO2 compensation point increased with salinity. Transpiration, dark respiration and photorespiration increased at low salinity but decreased at high salinity levels. In the field study, however, there were no significant differences in stomatal conductance and opening between saline and non-saline soils. Model predictions supported the results of the field measurements. Adaptation to salinity was reflected in an acclimatization of tree structure in the field study. There were both functioning and structural changes of seedlings in the greenhouse experiment
In terms of ecophysiological and morphological characteristics, E. Camaldulensis showed better salt tolerance than C. Quadragulare both in the greenhouse experiment and field study
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Genetic variation in the physiological characteristics and biomass accumulation of Acacia mangium Willd. was studied in both field and laboratory conditions. Variation in the growth characteristics, foliar nutrient concentration, phyllode anatomy and stomatal frequency was analysed in 16 different origins under field conditions in Central Thailand. Family variation and heritability of growth and flowering frequency were calculated using 20 open-pollinated families at the age of 28 months. The effect of environmental factors on diameter growth in different provenances is also discussed.
Under laboratory conditions, such physiological characteristics as transpiration rate, leaf conductance and leaf water potential were measured at varying soil moisture conditions. The responses of photosynthesis, photorespiration and dark respiration as well as the CO2 compensation point to temperature and irradiance were also investigated. All physiological characteristics indicated differences among provenances. An attempt was made to relate the results obtained in the laboratory to the growth performance in the field. Recommendations on provenance selection for the planting of A. mangium in Thailand are also given.
The PDF includes a summary in Finnish.
The dependence of the rates of photosynthesis and transpiration are studied on the environmental factors and on the control of the plant metabolism with Scots pine (Pinus sylvestris L.) in the natural environment. The importance of the different environmental factors to the photosynthetic production are evaluated. In modelling the dependence of the rates of photosynthesis and transpiration on the environmental factors and on the control processes, a dynamic system analysis approach is applied. Irradiance, temperature and water are explored during the annual cycle. Field measurements of the CO2 exchange and environmental factors over three years are used in the analysis of the rate of the photosynthesis. The effect of different environmental factors on photosynthetic production is evaluated by model estimations using weather data in a 20-year period.
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The problems caused by the temporal and spatial microvariation in irradiance during field measurements of photosynthesis are studied. It is concluded on the basis of variation analyses based on irradiance data measured in a Scots pine (Pinus sylvestris L.) stand that the microvariation should be measured by integrating it over the measurement time and space.
However, the curvlinearity of the light response of photosynthesis results in biased estimates when linear integration (mean irradiance) is used. The significance of the bias is examined using a simulation technique on irradiance material. Whether the actual integral of photosynthesis can be approximated with mathematical method is next studied. The method gave satisfactory results only for a low curvature response, but the latter method was applicable also to the high curvature response. However, both methods presuppose that the mean and variance are known. Measurement of the variance is based on integration of the second power.
A new method, where the nonlinearity problem is avoided, is presented to measure fluctuation of the irradiance. The method enables the shoot geometry to be taken into account and it is also applicable to transpiration studies.
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The report concludes a series of studies on the early development of young Scots pine (Pinus sylvestris L.) stands. The basis assumption made in the study series was that the within-stand light regime is the main driving force for total tree growth and its allocation of photosynthates for crown, stem and root growth. An individual tree growing in a stand under a varying light regime which is controlled by the stand structure, is the basic unit used in the study. The photosynthesis of an individual tree is determined by the light regime. The stand is formed from individual trees.
The model is applied in simulation of the growth and development of tree stands. Several computer runs representing various densities, height distributions and tree species mixtures were carried out. Potential application areas, properties of the model and future needs of investigations are discussed.
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The net photosynthetic rate per unit of foliage was studied in two-year old cuttings of Norway spruce (Picea abies (L.) H. Karst.), representing four clones, at varying temperature and soil moisture. The CO2 compensation point (Γ), photorespiration, dark respiration, and water balance were also investigated. All these characteristics indicated differences among the clones. A correlation between CO2 exchange and transpiration suggested that stomatal control determined at least a part of this variation during a favourable water balance. An inverse relationship existed between Γ and net photosynthetic rate, and the same curvilinear model explained this variation in unstressed as well as stressed plants at a given temperature. An increase in Γ seems to be a normal result of water stress, particularly at high temperature, indicating an increase in mesophyll resistance to CO2 diffusion. This result was in agreement with calculated values of mesophyll resistance. It also supported our earlier conclusions about the significance of mesophyll resistance during water stress.
In the experiment Scots pine (Pinus sylvestris L.) seedlings were transplanted out in the field. The effect of the treatments on gas metabolism and daily height increment were examined. The seedlings were 5-year old Scots pine plants growing in clay pots, covered with plastic bags. Transpiration and photosynthetic rates were monitored with open IRGA measuring system for a few days before being subjected to the treatments and for one month after. In addition, the daily amounts of transpired water and daily height increments were measured. A model for the potential rate of each metabolic process was constructed.
Planting and additional exposure had a strong and rather permanent effect on the self-regulation of the processes. This effect is very similar to that caused by water deficit. Exposure makes the disturbance more pronounced. Transpiration of the transplanted seedlings decreased in a few days after planting to less than half of the potential value and that of the exposed ones decreased to a quarter of the potential value. The daily amounts of photosynthesis decreased to half of the potential value. There was no recovery in photosynthesis during the whole monitoring period of four weeks. There was a slight recovery in transpiration about five weeks after transplanting.
Thus, the treatment probably generated stress conditions throughout the whole growing period, which is characterized by strong self-regulation of photosynthesis and transpiration, thus causing an essential decrease in the total amount of CO2 fixed. The photosynthesis was depressed especially at elevated temperatures after planting, as during water deficit. Planting and additional exposure did not produce any detectable changes in the dependence of the growth rate on temperature or in the effect of self-regulation on height growth. On the other hand, the level of growth was decreased as a result of planting out.
The PDF includes a summary in Finnish.