Diurnal changes in needle gas exchange in alpine Pinus pumila during snow-melting and summer seasons

Pinus pumila (Pallas) Regel. is a dominant dwarf tree in alpine regions of Japan. The possible factors limiting the net photosynthetic rate (P_n) of the needles of P. pumila were examined in the snow-melting (May) and the summer (August) seasons. In August, in situ maximum P_n was 20 mol kg^–1 needle s^–1 in the current-year needles and 25 mol kg^–1 needle s^–1 in the 1-year-old needles. Diurnal trends of P_n in August were positively related to fluctuations in photosynthetic photon flux density (PPFD) and no midday depression of P_n was found, indicating that a decrease in PPFD rather than an increase in needle-to-air vapor pressure deficit (W) might cause the reduction of P_n. Both stomatal conductance (g_s) and P_n were lower in May than in August. In May, P_n and g_s were almost zero in the morning, but gradually increased with decreasing W, reaching maximum P_n values (4 mol kg^–1 needle s^–1) and g_s (60 mmol kg^–1 needle s^–1) at 16.00 hours. The daytime P_n in May was positively related to g_s. Relative water content in the exposed needles above the snow in May was 83%, which was far above the lethal level. This indicates that the water flow from stems or soils to needles was enough to compensate for a small amount of water loss due to the low g_s in May, although the water supplied to needles would be impeded by the low temperatures. Thus, the reduced g_s in May would be important for avoiding needle desiccation, and would result in a reduced P_n.     

干旱和遮荫对马尾松幼苗生长和光合特性的影响

为探究马尾松对干旱和遮荫胁迫的生理响应规律和适应机制,以2年生马尾松幼苗为对象,设置对照(CK)、模拟干旱(DR)、遮荫(LL)以及干旱与遮荫的交互处理(DRLL)4种环境,研究干旱和遮荫对马尾松幼苗的生长和光合生理特性的影响。结果表明:(1)在干旱、遮荫和二者的交互处理下,马尾松幼苗的基径和株高增长量均显著减小,且二者的交互处理加重了干旱和遮荫单一处理下的减小趋势,二者交互作用的影响符合"相互作用理论"。(2)在干旱处理下,针叶长度和比叶面积减小,净光合速率、蒸腾速率、气孔导度和细胞间隙CO_2浓度显著降低,水分利用效率显著增加,光合色素含量基本不变。(3)在遮荫处理下,针叶长度和比叶面积增大,净光合速率、蒸腾速率、气孔导度显著降低,光合色素含量显著增加,水分利用效率和细胞间隙CO_2浓度基本未发生改变。(4)二者的交互处理下,针叶长度、比叶面积减小,净光合速率、蒸腾速率和气孔导度显著降低,且降低程度大于干旱单一处理,光合色素含量显著增加,但其增加程度小于遮荫单一处理。说明干旱和遮荫均能抑制马尾松植株的生长,但其光合生理特性在干旱和遮荫胁迫下分别表现出不同的响应特征。在干旱处理下,通过牺牲针叶长度和比叶面积、调节气孔导度达到保水目的来适应干旱环境;在遮荫处理下,通过增大针叶长度、比叶面积和光合色素含量来响应遮荫环境。在二者的交互处理下,干旱胁迫导致马尾松针叶长度、比叶面积和气孔导度的减小趋势不仅未因遮荫环境而得到缓解,而且还削弱了对光合色素的合成能力,因此,二者的交互作用会进一步加重对马尾松生长的抑制。     

The effect of vapor pressure on stomatal control of gas exchange in Douglas fir (Pseudotsuga menziesii) saplings

Summary Increasing leaf to air vapor pressure deficit (VPD) caused reductions in stomatal conductance of both current year and previous season needles of Pseudotsuga menziesii saplings. The stomata of current year needles were found to be more responsive to changes in VPD than those of previous season needles. The reductions in stomatal conductance of current year needles were not associated with decreases in xylem pressure potential. In fact, the reductions in stomatal conductance of current year needles were sometimes sufficient to reduce transpiration and thus raise xylem pressure potential even though VPD was increasing. There was a decline in stomatal responsiveness to VPD in current year needles between early and late summer. Pressure-volume curves determined for different age needles at different times of the year suggested that differences and changes in stomatal responsiveness to VPD may have been caused in part by differences and changes in needle water potential components. Hexane washes of current year needles during the late summer succeeded in partially restoring their VPD sensitivity, suggesting that changes in the water permeability of the external cuticle during needle maturation may also have played a role in causing the summer decline in VPD responsiveness.In both current and previous year needles VPD-induced changes in stomatal conductance had a greater relative effect on transpiration (q _w) than on net photosynthesis (Ph_N). In maturing needles the ratio of the sensitivities of transpiration and net photosynthesis to changes in stomatal conductance, (q _w/g _s)/Ph_N/g _s), remained nearly constant as VPD was varied. This provides experimental support for a recent hypothesis that stomata respond to environmental fluctuations in such a manner as to maintain the above ratio constant, which optimizes CO₂ uptake with respect to water loss.     

Carbon isotope discrimination and stomatal responses of mature Pinus sylvestris L. trees exposed in situ for three years to elevated CO2 and temperature

The Climate Change Experiment (CLIMEX) is a unique large scale facility in which an entire undisturbed catchment of boreal vegetation has been exposed to elevated CO₂ (560 ppm) and temperature (+3°C summer, +5°C winter) for the past three years with all the soil-plant-atmosphere linkages intact. Here, carbon isotope composition and stomatal density have been analysed from sequential year classes of needles of mature Scots pine trees (Pinus sylvestris L.) to investigate the response of time-integrated water-use efficiency (UWE) and stomatal density to CO₂ enrichment and climate change. Carbon isotope discrimination decreased and WUE increased in cohorts of needles developing under increased CO₂ and temperature, compared to needles on the same trees developing in pretreatment years. Mid-season instantaneous gas exchange, measured on the same trees for the past four years, indicated that these responses resulted from higher needle photosynthetic rates and reduced stomatal conductance. Needles of P. sylvestris developing under increased CO₂ and temperature had consistently lower stomatal densities than their ambient grown counterparts on the same trees. The stomatal density of P. sylvestris needles was inversely correlated with δ¹³C-derived WUE, implying some effect of this morphological response on leaf gas exchange. Future atmospheric CO₂ and temperature increases are therefore likely to improve the water economy of P. sylvestris, at least at the scale of individual needles, by affecting stomatal density and gas exchange processes.     

Why does photosynthesis decrease with needle age in Pinus pinaster?

Rates of photosynthesis vary with foliage age and typically decline from full-leaf expansion until senescence occurs. This age-related decline in photosynthesis is especially important in species that retain foliage for several years, yet it is not known whether the internal conductance to CO₂ movement (g _i) plays any role. More generally, g _i has been measured in only a few conifers and has never been measured in leaves or needles older than 1 year. The effect of ageing on g _i was investigated in Pinus pinaster, a species that retains needle for 4 or more years. Measurements were made in autumn when trees were not water limited and after leaf expansion was complete. Rates of net photosynthesis decreased with needle age, from 8 μmol m⁻² s⁻¹ in fully expanded current-year needles to 4.4 μmol m⁻² s⁻¹ in 3-year-old needles. The relative limitation due to internal conductance (0.24–0.35 out of 1) was in all cases larger than that due to stomatal conductance (0.13–0.19 out of 1). Internal conductance and stomatal conductance approximately scaled with rates of photosynthesis. Hence, there was no difference among year-classes in the relative limitations posed by internal and stomatal conductance or evidence that they cause the age-related decline in photosynthesis. There was little evidence that the age-related decline in photosynthesis was due to decreases in contents of N or Rubisco. The decrease in rates of photosynthesis from current-year to older needles was instead related to a twofold decrease in rates of photosynthesis per unit nitrogen and V _cmax/Rubisco (i.e., in vivo specific activity).     

Needle CO2 exchange, structure and defense traits in relation to needle age in Pinus heldreichii Christ – a relict of Tertiary flora

 Pinus heldreichii Christ is a long-lived, slow-growing Tertiary relict from the Balkans. In this study we evaluated the physiological characteristics of eight needle-age classes of P. heldreichii grown at the Arboretum of the Institute of Dendrology in Kórnik, Poland. At the end of the growing season, current-year foliage had the highest rates of mass-based light-saturated net photosynthesis (A_sat) of 33.5 nmol CO₂· g^–1· s^–1. A_sat decreased with needle age, but older needle classes retained from approximately 62 to 26% of the current needles’ rate. The relationship between leaf N and chlorophyll a concentration among all needle-age classes was highly significant (r = 0.96, P = 0.0006). The variation in A_sat of 1- to 7-year-old needles was linearly related to needle N concentration (r = 0.98, P = 0.0001). Needle dark respiration rates among these needle age classes ranged from 0.8 to 2.2 nmol · g^–1· s^–1 and decreased with needle age and nitrogen concentration. Total phenols and glucose concentrations increased linearly with needle age. A similar pattern was observed in acid buffering capacity and the pH of tissue homogenates. The water content ranged from 62% for the current needles to 51% for the 6-year-old needles. Greater investment in leaf structural tissue and increased chemical defense is associated with higher structural cost of older needles and may reduce their photosynthetic activity. Significant declines in water and nitrogen content with needle age and an increase in content of phenolics is most likely a defense adaptation of P. heldreichii related to the species’ long-lived leaves. Received: 8 January 1997 / Accepted: 4 March 1997     

Surface areas,lengths and volumes of Picea abies (L.) Karst. needles: determination,biological variability and effect of environmental factors

Summary A method for the rapid determination of the lengths and surface areas of very large samples of needles of Picea abies (L.) Karst. using a computer-aided image analysis system was developed. Two independent methods for measuring non-destructively the volumes of individual needles and of all needles attached to a twig were devised. The surface areas and lengths of about 38000 needles sampled from the three youngest needle age-classes (1986, 1985, 1984) of 48 trees approximately 130 years old at four sites in the Fichtelgebirge mountains (N. E. Bavaria, FRG) were measured. The frequency distributions of lengths and areas for each site and age-class are given. Variability of needle size was fairly large. Even though the sites differed in climate, soil, and air pollution levels no consistent effect of these factors on needle size could be detected. Needle lengths and surface areas did not correlate with either the total chlorophyll content of the needles or the degree of crown thinning. The needle surface area (in mm²) of fully developed P. abies needles can be estimated by the empirical equation surface area = 4.440 x needle length -24.8 (r = 0.937), and the needle volume (in mm³) by needle volume = 0.208 x projected needle area ^1.353 (r = 0.969).     

Effects of CO2 concentration and irradiance on the stomatal behaviour of maize, barley and sunflower plants in the field

Abstract Maize, barley and sunflower plants were grown in the field, well supplied with water and nutrients. During growth, net CO₂ exchange and transpiration of the crops at varying ambient CO₂ concentrations and irradiance were determined by infra-red gas analysis. In maize the net photosynthetic rate (P_n) was linearly related to the irradiance (I) and independent of the ambient CO₂ concentration (C_a). The transpiration rate (E_T) was also linearly related to I but decreased strongly with increasing C_a. In sunflower and barley P_n increased and E_T decreased with increasing C_a. A mean stomatal conductance and intercellular CO₂ concentration (C_i) were calculated. In all three species the internal CO₂ concentration was independent of the irradiance. In maize it was also independent of C_a, but in sunflower and barley C_i was proportional to C_a with a ratio of 0.6. It is concluded that differences in stomatal behaviour are only partly species-specific and depend mainly on growing conditions. The importance of stomatal regulation for crop growth under conditions of water shortage and CO₂ depletion is discussed.     

Effects of long-term open-air exposure to fluoride,nitrogen compounds and SO2 on visible symptoms,pollutant accumulation and ultrastructure of Scots pine and Norway spruce seedlings

 Effects of SO₂, aqueous fluoride (NaF) and a solution of nitrogen compounds (NH₄NO₃) on the visible symptoms, pollutant accumulation and ultrastructure of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] seedlings were studied in an open-air experiment lasting for 3 consecutive years. Visible injury symptoms were most pronounced in combination exposures and whenever F was applied. Visible symptoms correlated well with needle pollutant concentrations. Exposure to NaF increased needle F contents particularly when F was applied with SO₂ or NH₄NO₃. This suggests that a reduction in N or SO₂ emissions, in F polluted areas, could improve the condition of conifers via decreased accumulation of phytotoxic F in the needles. Norway spruce needles accumulated 2 – 10 times as much S and F as those of Scots pine. Microscopic observations showed various changes in the needle mesophyll cell ultrastructure. In both species, exposure to SO₂ increased significantly the amount of cytoplasmic vacuoles, suggesting detoxification of excess sulphate or low pH. F treatments resulted in a significant enlargement of plastoglobuli in Scots pine and a darkening of plastoglobuli in Norway spruce. All exposures enhanced the accumulation of lipid bodies. An increased portion of translucent plastoglobuli was most pronounced in N treatments. Many of the ultrastructural changes and visible symptoms appeared only as number of years exposed increased, indicating that long-term experiments are needed. Both visible symptoms and ultrastructural changes pointed to the more pronounced sensitivity of Norway spruce compared to Scots pine. Ultrastructural results mostly supported earlier qualitative observations of F, N and SO₂ effects on needle mesophyll cell ultrastructure. However, no reduction of thylakoids in SO₂ containing exposure or curling of thylakoids in F exposure could be detected in the present study. Received: 5 December 1994 / Accepted: 28 April 1995     

Soil type affects nitrogen conservation in foliage of small Pinus sylvestris L. trees

Nutrient conservation in plants and soil fertility may be intricately linked. We studied nitrogen conservation in small Scots pine (Pinus sylvestrisL.) trees growing in stands on organogenic Dystric Histosols and on mineral Podzols. Nitrogen-resorption efficiency (NRE) and proficiency (NRP) of senescent needles, and mean residence time of nitrogen (MRT) were studied in relation to needle surface area, needle longevity, and leaf mass per area (LMA). Trees on Podzols had higher nitrogen concentration in green needles than the trees on Dystric Histosols, but the nitrogen concentration of yellowing needles was similar for trees on both soil types. NRE averaged 65±3.5% (mean±SD) and 56±7.2% for the trees on Podzols and Dystric Histosols, respectively. Neither NRP (0.44±0.05% and 0.35±0.07%, respectively) nor MRT (8.4±2.3 and 6.1±1.2 years) differed significantly between the stands on the two soil types. Mean needle surface area was significantly smaller in trees on Dystric Histosols (76±29 mm²) than on Podzols (131±38 mm²), whereas needle longevity varied between 2 and 4 years independently of the soil type. Trees invested, on average, the same amount of dry matter per unit of needle area on both soil types. Growth of trees, measured as increment of shoot length, was more restricted on Dystric Histosols (55±18 mm yr^–1) than on Podzols (184±44 mm yr^–1). The results of the correlation analysis applied to pooled data were inconsistent with the relations between traits of stress resistance syndrome observed in inter-specific comparisons. The study indicated that Scots pine trees relocated nitrogen from senescent foliage more efficiently on mineral Podzols than on organogenic Dystric Histosols, but the minimum nitrogen concentration of needles appeared to be similar on both soil types.     

Estimating the uptake of traffic-derived NO2 from 15N abundance in Norway spruce needles

The ¹⁵N ratio of nitrogen oxides (NO_x) emitted from vehicles, measured in the air adjacent to a highway in the Swiss Middle Land, was very high [δ¹⁵N(NO₂) = +5.7‰]. This high ¹⁵N abundance was used to estimate long-term NO₂ dry deposition into a forest ecosystem by measuring δ¹⁵N in the needles and the soil of potted and autochthonous spruce trees [Picea abies (L.) Karst] exposed to NO₂ in a transect orthogonal to the highway. δ¹⁵N in the current-year needles of potted trees was 2.0‰ higher than that of the control after 4 months of exposure close to the highway, suggesting a 25% contribution to the N-nutrition of these needles. Needle fall into the pots was prevented by grids placed above the soil, while the continuous decomposition of needle litter below the autochthonous trees over previous years has increased δ¹⁵N values in the soil, resulting in parallel gradients of δ¹⁵N in soil and needles with distance from the highway. Estimates of NO₂ uptake into needles obtained from the δ¹⁵N data were significantly correlated with the inputs calculated with a shoot gas exchange model based on a parameterisation widely used in deposition modelling. Therefore, we provide an indication of estimated N inputs to forest ecosystems via dry deposition of NO₂ at the receptor level under field conditions. Received: 7 November 1997 / Accepted: 16 September 1998     

Genotypic variation for drought tolerance in cotton (Gossypium hirsutum L.): Leaf gas exchange and productivity

Although water-limited environments are detrimental to cotton growth and productivity worldwide, identification of cotton (Gossypium hirsutum L.) genotypes that are less sensitive to drought may improve productivity in drought prone areas. The objective of the study was to assess genotypic variation for drought tolerance in cotton varieties using physiological attributes as selection criteria, and to determine the relationship of physiological attributes with productivity traits. The association of target physiological traits for drought tolerance (photosynthetic rate (P_n), stomatal conductance (g_s), and transpiration rate (E)) with productivity traits under well-watered (W₁) and water-limited (W₂) regimes was analyzed using 32 public cotton cultivars/bred lines in two field experiments conducted during the normal cotton growing seasons 2003 and 2004. Seed cotton yield (SCY) and biological yield (BY) were markedly affected under W₂ regime in all cultivars except the outstanding performance of CIM-1100 and RH-510 proving their superiority to other cultivars in drought tolerance. Conversely, FH-901, FH-634, and FH-2000 were high yielding under W₁ regime; however, exhibited a sharp decline in yield under W₂ regime. A positive correlation between SCY and BY under water stress (r=0.44 in 2003; r=0.69 in 2004) indicates that BY is also a primary determinant of SCY under water stress and genetic improvement of BY under water-limited environment may also improve SCY. P_n, g_s, and E were significantly reduced by water stress. Substantial genotypic variation for gas exchange attributes existed among the cotton cultivars. A positive association (P<0.01) was observed between g_s and E under both regimes in both years indicating the prevalence of stomatal control of transpiration. The positive association (P<0.01) between P_n and g_s in both years in W₂ regime indicates also a major role of stomatal effects in regulating leaf photosynthesis under water-limited conditions. P_n was significantly correlated with SCY (P<0.01) and BY (P<0.05 in 2003; P<0.01 in 2004) in W₂ regime; however, the level of these associations was not significant in W₁ regime. These findings demonstrate that association of P_n with productivity is effective under water-limited environment and may be useful as a selection criterion in breeding programs with the objective of improving drought tolerance and SCY under water-limited environments. Moreover, association between SCY and BY under water stress suggests that genetic improvement of BY under water stress may also improve SCY.     

Photosynthesis and transpiration ofPinus radiata D. Don under natural conditions in a forest stand

Summary Gas-exchange ofPinus radiata foliage was measured with climatised cuvettes under natural light in the sun-crown of 8 m tall trees in a forest stand. Measurement began during a period of drought (_WS –8.2 bar, _We –10.5 bar) and continued after elimination of soil moisture-deficit by watering (_WS –0.5 bar, _We –5.5 bar). Soil and air moisture-deficits severely restricted gas-exchange. Watering resulted in an immediate decline in stomatal resistance (r _s ) and an increase in net photosynthesis (P _N ) of 13%. A slower progressive gas-exchange recovery occurred additionally during the 10 days after watering leading to a further decline inr _s to 3 s cm^-1 and an ultimate increase inP _N of 38% when measured under comparable conditions at 8 mb v.p.d. Radiata pine had a high photosynthetic capacity with a measured maximumP _N of 10.2 mg CO₂ dm^-2 h^-1 total needle surface (11.4 mg CO₂ g^-1 DM h^-1).Optimum temperature forP _N in March (late summer) occurred at ca. 18°C. Rate ofP _N was 95% saturated at irradiance of 900 E m^-2 s^-1 and 50% saturated at only 270 E m^-2 s^-1. Radiata pine needles responded directly to changes in atmospheric humidity by adjusting their stomatal diffusive resistance. As a result, between 8 and 18 mb v.p.d.P _N declined by 2.3% mb^-1 increase.     

Seasonal changes in needle water content and needle ABA concentration of Japanese red pine, Pinus densiflora, in declining forests on Mt. Gokurakuji, Hiroshima prefecture, Japan

To evaluate the effects of air pollution on the decline of Pinus densiflora forests, various research has been conducted around Mt. Gokurakuji (34° 23N, 132° 19E, 693 m a.s.l.) north of the Seto Inland Sea, west Japan. To investigate the mechanisms responsible for decreases in photosynthesis (Pn) and stomatal conductance (gl), ¹³C of needles and seasonal changes in the water content (WC) and abscisic acid concentration (ABA) of needles were measured in various stands. The ¹³C values were less negative in declining stands and younger needles. ABA and WC were not correlated with each other. WC decreased consistently with needle age while the ABA showed a minimum in August and a smaller content in older needles. Monthly precipitation and the daily maximum vapor pressure were not correlated with ABA and WC. In declining stands, WC and ABA tended to be higher and lower, respectively, than in nondeclining stands. These results suggest that the trees in declining stands received less water stress than those in nondeclining stands and the differences in gl and ¹³C are not caused by the difference in water stress. The possibilities of the effects of air pollution and the infection of pine-wood nematode on the physiological decline on the pine needles are discussed.     

Water relations of Picea abies. II. Determination of the apoplastic water content and other water relations parameters of needles by means of capillary microcryoscopy and the pressure-volume

Using the pressure volume analysis (PV analysis) on the shoots of Norway spruce (Picea abies [L.] Karst.) and the here presented capillary microcryoscopy of the needle press sap of the same shoots, it was possible to determine the amount of apoplastic water in the needles (W_an) as well as in the defoliated shoots (W_as). Additionally, the bulk osmotic pressure at full water saturation in the symplast of the needles and defoliated shoots (π_on and π_os) was determined. The dependence of the bulk-averaged turgor pressure (P_t) on the water content and the relationship between the bulk modulus of elasticity of the needles (?_n) and the bulk-averaged needle turgor pressure (P_tn) was shown with help of the PV analysis on the whole shoots and defoliated shoots. The study was conducted at the end of the vegetation period in 1987 and during winter 1988. The proportion of W_an in the total needle water content (W_tn) was 14% in September 1987 and 12.5% in March 1988. The respective percentage of W_as in W_ts were 27% and 25%. The amount of apoplastic water depended on the ratio of the dry weight of defoliated shoots to the dry weight of the whole shoots. A standard mean value for the amount of W_an in the total water content of the shoots (W_t) was therefore not possible. The bulk osmotic pressure at full water saturation in the needle symplasts was –1.9 MPa in September 1987 and –2.2 MPa in winter 1988. The respective values of the bulk osmotic pressures in the symplast of the defoliated shoots (π_os) were –1.5 MPa and –1.7 MPa. Thus π_on was 0.1 MPa lower and π_os 0.3–0.4 MPa higher than the average osmotic pressure during full water saturation in the symplast of the whole shoots (π_o). The relation between bulk-averaged turgor pressure and water content showed that during water loss P_tn dropped more rapidly than the turgor pressure of defoliated shoots (P_ts). Consequently the needles were less elastic than the defoliated shoots. The turgor values of whole shoots followed an intemediate course between P_tn and P_ts. The flat course of P_ts seems to be the main reason for the often observed “plateau” of ψ-isotherms of whole shoots near full turgor.     

Stomatal SO2 uptake and sulfate accumulation in needles of Norway spruce stands (Picea abies) in Central Europe

Monthly uptake rates and the annual deposition of gaseous SO₂ via the stomata of six Norway spruce canopies (Picea abies (L.) Karst.) in Germany (Königstein im Taunus, Witzenhausen, Grebenau, Frankenberg, Spessart, Fürth im Odenwald) were calculated (i) from statistical response functions of stomatal aperture depending on meteorological data, and (ii) from the synchronously measured SO₂ immission at these stands. The stomatal response functions had been derived on the basis of thorough stomatal water conductance measurements in the field. Calculations of the SO₂ conductance of spruce twigs and SO₂ uptake rates via stomata need continuously measured complete data sets of the (i) light intensity, (ii) air temperature, (iii) air humidity and (iv) SO₂ concentration in spruce forests from all the year. These data were recorded half hourly in different German spruce forests. The apparent needle water vapour pressure difference and transpiration rates were calculated from meteorological data. Additional use of canopy through flow data in dry years allowed the estimation of the mean stomatal conductance for H₂O and SO₂ of whole spruce canopies. The annual SO₂ uptake of a mean unit needle surface in spruce forests was 32% of the SO₂ uptake rate of exposed needles at the top of spruce crowns. There is significant SO₂ uptake all the year. The mean SO₂ dose at all sites and years received through the stomata was (0.25±0.07) mol SO₂ m^-2 (total needle surface) (nPa Pa^-1)^-1 (annual mean of SO₂ immission; 1 nPa (SO₂) Pa^-1 (air) = 1 ppb) day^-1 (vegetation period per year). Comparison of calculated SO₂ uptake rates into needles with measured SO₄ ^2- accumulation rates in needles from the mentioned sites and additionally from Würzburg, Schneeberg (Fichtelgebirge) and from three sites in the eastern Erzgebirge (Höckendorf, Kahleberg, Oberbärenburg) revealed that oxidative SO₂ detoxification (SO₄ ^2- formation) dominates only at sites with high SO₂ immission and short vegetation periods. Under these conditions 70 to 90% of the annual stomatal SO₂ uptake is detoxified via SO₄ ^2- accumulation in needles. Cations are needed for neutralization of accumulating SO₄ ^2- which are inavailable to support growth. Thus, SO₂ induces a dominant and competitive additional nutrient cation demand, cation deficiency symptoms and enhanced needle loss (spruce decline symptoms) mainly at sites, where the ratio R=(SO₂ immission): (length of the vegetation period) is higher than R=0.07 nPa Pa^-1 day^-1. Correlation analysis of the relative needle loss versus the SO₂-dependent SO₄ ^2- formation rate revealed a significant increase of needle loss at the 98% level (Student). At sites with small SO₂ immission and long vegetation periods (R<0.07 nPa Pa^-1 day^-1) reductive SO₂ detoxification via growth (and/or phloem export of SO₄ ^2-) is not kinetically overburdened. Under these conditions only 30% of the annual SO₂ uptake is detoxified via SO₄ ^2- formation and spruce decline is small or absent. On the basis of the critical value R0.07 nPa Pa^-1 day^-1 recommended SO₂ immission limits can be deduced on a mere ecophysiological basis. These deduced values are close to the proposed SO₂ immission limits of the IUFRO, WHO and the UNECE.     

Light microscopic structure of needles of Scots pine (Pinus sylvestris L.) in relation to air pollution and needle element concentrations in S.E. Finland and the Karelian Isthmus, N.W. Russia

 The mesophyll structure of current and second year needles of mature Scots pines (Pinus sylvestris L.), growing on a survey line from the Karelian Isthmus (N.W. Leningrad region, Russia) to S.E. Finland, was studied at the light microscopic level in 1992 and 1994. The area is affected by sulphur dioxide and nitrogen oxide emissions from Russia (200  000–285 000 ton/year SO₂ and about 100  000 ton/year NO_X), Estonia (140  000–320  000 ton/year SO₂ and 30  000–50  000 ton/year NO_X) and Finland (6000–9000 ton/year SO₂ and 23 000–25 000 ton/year NO_X) and by unknown amounts of neutralizing compounds from Estonia. Mean concentrations of SO₂ were between 1.0 and 1.5 μg m^–3 and those of ozone between 52 and 61 μg m^–3 during growing seasons of 1991–1994. The concentrations tended to be lower in 1993/94 than in 1991/92. The results showed cellular changes related to exposure to sulphur dioxide, ozone and acid wet deposition. The percentage of cells with injuries related to acid deposition was higher in 1994 than in 1992, possibly due to higher reduction of neutralizing than of acidifying compounds in the main emission areas, while the percentage of cells with injuries related to sulphur dioxide correlated positively with needle sulphur concentrations and showed a declining trend between Karelian Isthmus and S.E. Finland in the needles sampled in 1992. These changes and those related to ozone were considerably less numerous in 1994 than in 1992, possibly on account of the lower concentrations of SO₂ and ozone in 1993/94 than in 1991/92, especially in early summer, the time of the most sensitive phase of needle growth. In addition, periods of lower wet deposition in summer 1992, and considerably lower temperatures in autumn/winter 1992 than in 1994 may have enhanced the injuries in the needles collected in October/December 1992, relative to those collected in 1994. The results of the microscopic work agreed well with the simultaneous chemical measurements, pointing to the usefulness of cellular analysis for diagnostic work in the field. Received: 26 September 1997 / Accepted: 24 November 1997     

Physiological effects of a long term exposure to low concentrations of NH3, NO2 and SO2 on Douglas fir (Pseudotsuga menziesii)

The above-ground parts of two years old seedlings of Douglas fir (Pseudotsuga menziesii) were exposed to filtered air, NH₃, NO₂₊, SO₂ (66, 96 and 95 μg m^?3, respectively), to a mixture of NO₂+NH₃ (55 + 82 μg m^?3) or SO₂+NO₂ (128 + 129 μg m^?3), for 8 months in fumigation chambers. Both chlorophyll fluorescence and gas exchange measurements were carried out on shoots which had sprouted at the beginning of the exposure period. The chlorophyll fluorescence measurements were performed after 3 and 5 months of exposure (average shoot age 70 and 140 days, respectively). Light response curves of electron transport rate (J) were determined, in which J was deduced from chlorophyll fluorescence. In addition, light response curves of net CO₂ assimilation were determined after 5 months of exposure. After 3 months of exposure (average shoot age 70 days) all exposure treatments showed a lower maximum electron transport rate (J_max) as compared to the control shoots (filtered air). A large reduction (45%) was observed for shoots exposed to SO₂+NO₂. During the exposure period between 3 and 5 months (average shoot age 70 and 140 days, respectively) a decrease of J_max was observed for all treatments. J_max had further declined some time after termination of the exposure, when average shoot age was 310 days. Shoots exposed to SO₂ and SO₂+NO₂ also showed a reduction in maximum net CO₂ assimilation (P_max) as compared to the control shoots. However, shoots exposed to NO₂ showed no reduction and even a higher P_max was observed for shoots exposed to NH₃ or NO₂+NH₃. Needles of these treatments also showed a higher chlorophyll content which might explain the contradictory results obtained for these treatments: the increased amount of photosynthetic units counteracts the reduction in J_max and consequently no reduction in P_max is measured. Shoots exposed to SO₂ and SO₂+NO₂ also showed a reduction in maximum stomatal conductance (g_s). However, the stomatal opening was larger than could be expected on basis of their (maximum) CO₂ assimilation rate. Consequently, water use efficiency of these shoots was lower than that of the control shoots. Also shoots exposed to NO₂ had a lower water use efficiency due to a significantly higher maximum g_s. Shoots exposed to NH₃ showed a high transpiration rate in the dark, indicating imperfect stomatal closure.     

Physiological responses to salinity in Silver buffaloberry (<Emphasis Type="Italic">Shepherdia argentea</Emphasis>) introduced to Qinghai high-cold and saline area,China

Since 2002, Silver buffaloberry (Shepherdia argentea) has been introduced from North America in order to improve the fragile ecological environment in western China. To elucidate the salt-resistance mechanism of S. argentea, we conducted a test with two-year-old seedlings subjected to 0, 200, 400, and 600 mM NaCl solutions for 30 d. The results showed that significant salt-induced suppression of plant fresh mass (FM) and stem height of S. argentea seedlings occurred only at the highest salinity level (600 mM). Leaf number, plant dry mass (DM), and chlorophyll (Chl) content declined markedly at both 400 and 600 mM. Leaf area (LA) and leaf water potential (Ψ_w) continuously declined with the increase of salinity. There was also a progressive and evident decrease in net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) with the increase of salinity and time. The correlation analysis indicated that P _N was positively correlated with g _s at all salinity levels while correlated with intercellular CO₂ concentration (C _i) only at moderate salinity levels (<600 mM). Based on the initial slope of the P _N/C _i curves, the estimated carboxylation efficiency (CE) was strongly inhibited at 600 mM. We confirm that S. argentea is highly tolerant to salinity. Moreover, our results show that at moderate salinity levels, salt-induced inhibition of photosynthesis is mainly attributed to the stomatal efficient closure predetermined by a low water potential in leaves; while at the high salinity levels, the inhibition is mainly due to the suppression of chloroplast capacity to fix CO₂ caused by the serious decline in both CE and Chl contents.     

Nitrogen allocation and carbon isotope fractionation in relation to intercepted radiation and position in a young Pinus radiata D. Don tree

The three dimensional distribution of intercepted radiation, intercellular CO₂ concentration (C_i) and late summer needle nitrogen (N) concentration were determined at the tips of all 54 branches in a 6·2-m-tall Pinus radiata D. Don tree growing in a New Zealand plantation. Measurements included above- and below-canopy irradiance, leaf stable carbon isotopic composition (δ¹³C) and tree canopy architecture. The radiation absorption component of the model, MAESTRO, was tested on site and then used to determine the branch tip distribution of intercepted radiation. We hypothesized that in branch tip needles: (i) the allocation of nitrogen and other nutrients would be closely associated with the distribution of intercepted radiation, reflecting carbon gain optimization theory, and (ii) C_i would predominantly reflect changes in photosynthetic rate (A) rather than stomatal conductance (g_s), indicating that the increase in A for a given increase in N concentration was larger than the corresponding increase in g_s. Needle nitrogen concentration was poorly related to intercepted radiation, regardless of the period over which the latter was calculated. At a given height, there was a large azimuthal variation in intercepted radiation but N concentration was remarkably uniform around the tree canopy. There was, however, a linear and positive correspondence between N concentration and δ¹³C and needle height above ground (r² = 0·73 and 0·68, respectively). The very strong linear correspondence between N concentration and C_i (r² = 0·71) was interpreted, using gas exchange measurements, as supporting our second hypothesis. Recognizing the strong apical control in P. radiata and possible effects of leaf nitrogen storage in an evergreen species, we propose that the tree leader must have constituted a very strong carbon sink throughout the growing season, and that the proximity of branch tip needles to the leader affected their photosynthetic capacity and nutrient concentration, independent of intercepted radiation. This implies an integrated internal determination of resource allocation within the tree and challenges the current convention that resources are optimally distributed according to the profile of intercepted radiation.