The influence of elevated CO2 and O3 on fine roots and mycorrhizas of naturally growing young Scots pine trees during three exposure years

Young Scots pine trees naturally established at a pine heath were exposed to two concentrations of CO₂ (ambient and doubled ambient) and two O₃ regimes (ambient and doubled ambient) and their combination in open-top field chambers during growing seasons 1994, 1995 and 1996 (late May to 15 September). Filtered ozone treatment and chamberless control trees were also included in the treatment comparisons. Root ingrowth cores were inserted to the undisturbed soil below the branch projection of each tree at the beginning of the fumigation period in 1994 and were harvested at the end of the fumigation periods in 1995 and 1996. Root biomasses were determined from different soil layers in the ingrowth cores, and the infection levels of different mycorrhizal types were calculated. Elevated O₃ and CO₂ did not have significant effects on the biomass production of Scots pine coarse (Ø > 2 mm) or fine roots (Ø < 2 mm) and roots of grasses and dwarf shrubs. Elevated O₃ caused a transient stimulation, observable in 1995, in the proportion of tuber-like mycorrhizas, total mycorrhizas and total short roots but this stimulation disappeared during the last study year. Elevated CO₂ did not enhance carbon allocation to root growth or mycorrhiza formation, although a diminishing trend in the mycorrhiza formation was observed. In the combination treatment increased CO₂ inhibited the transient stimulating effect of ozone, and a significant increase of old mycorrhizas was observed. Our conclusion is that doubled CO₂ is not able to increase carbon allocation to growth of fine roots or mycorrhizas in nutrient poor forest sites and realistically elevated ozone does not cause a measurable limitation to roots within a period of three exposure years.     

Soil CO2 efflux of two silver birch clones exposed to elevated CO2 and O3 levels during three growing seasons

In the present open‐top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO₂) and ozone (O₃), singly and in combination, and soil CO₂ efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O₃ caused soil CO₂ efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO₂ was dependent on the genotype, as the soil CO₂ efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO₂ treatments. Like the O₃ impact, this clonal difference in soil respiration response to CO₂ increased as the experiment progressed. Although the O₃ impact did not differ significantly between clones, a significant time × clone × CO₂× O₃ interaction revealed that the O₃‐induced stimulation of soil respiration was counteracted by elevated CO₂ in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO₂ and O₃ in combination was almost constantly additive during the 3‐year experiment. Altogether, the root or above‐ground biomass results were only partly parallel with the observed soil CO₂ efflux responses. In conclusion, our data show that O₃ impacts may appear first in the below‐ground processes and that relatively long‐term O₃ exposure had a cumulative effect on soil CO₂ efflux. Although the soil respiration response to elevated CO₂ depended on the tree genotype as a result of which the O₃ stress response might vary considerably within a single tree species under elevated CO₂, the present experiment nonetheless indicates that O₃ stress is a significant factor affecting the carbon cycling in northern forest ecosystems.     

Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings

In this study, we investigated the impact of elevated atmospheric CO₂ (ambient + 350 μmol mol^–1) on fine root production and respiration in Scots pine (Pinus sylvestris L.) seedlings. After six months exposure to elevated CO₂, root production measured by root in-growth bags, showed significant increases in mean total root length and biomass, which were more than 100% greater compared to the ambient treatment. This increased root length may have lead to a more intensive soil exploration. Chemical analysis of the roots showed that the roots in the elevated treatment accumulated more starch and had a lower C/N-ratio. Specific root respiration rates were significantly higher in the elevated treatment and this was probably attributed to increased nitrogen concentrations in the roots. Rhizospheric respiration and soil CO₂ efflux were also enhanced in the elevated treatment. These results clearly indicate that under elevated atmospheric CO₂ root production and development in Scots pine seedlings is altered and respiratory carbon losses through the root system are increased.     

Effects of elevated atmospheric CO2 on net ecosystem CO2 exchange of a scrub–oak ecosystem

We report the results of a 2‐year study of effects of the elevated (current ambient plus 350 μmol CO₂ mol^?1) atmospheric CO₂ concentration (C_a) on net ecosystem CO₂ exchange (NEE) of a scrub–oak ecosystem. The measurements were made in open‐top chambers (OTCs) modified to function as open gas‐exchange systems. The OTCs enclosed samples of the ecosystem (ca. 10 m² surface area) that had regenerated after a fire, 5 years before, in either current ambient or elevated C_a. Throughout the study, elevated C_a increased maximum NEE (NEE_max) and the apparent quantum yield of the NEE (φ_NEE) during the photoperiod. The magnitude of the stimulation of NEE_max, expressed per unit ground area, was seasonal, rising from 50% in the winter to 180% in the summer. The key to this stimulation was effects of elevated C_a, and their interaction with the seasonal changes in the environment, on ecosystem leaf area index, photosynthesis and respiration. The separation of these factors was difficult. When expressed per unit leaf area the stimulation of the NEE_max ranged from 7% to 60%, with the increase being dependent on increasing soil water content (W_soil). At night, the CO₂ effluxes from the ecosystem (NEE_night) were on an average 39% higher in elevated C_a. However, the increase varied between 6% and 64%, and had no clear seasonality. The partitioning of NEE_night into its belowground (R_below) and aboveground (R_above) components was carried out in the winter only. A 35% and 27% stimulation of NEE_night in December 1999 and 2000, respectively, was largely due to a 26% and 28% stimulation of R_below in the respective periods, because R_below constituted ca. 87% of NEE_night. The 37% and 42% stimulation of R_above in December 1999 and 2000, respectively, was less than the 65% and 80% stimulation of the aboveground biomass by elevated C_a at these times. An increase in the relative amount of the aboveground biomass in woody tissue, combined with a decrease in the specific rate of stem respiration of the dominant species Quercus myrtifolia in elevated C_a, was responsible for this effect. Throughout this study, elevated C_a had a greater effect on carbon uptake than on carbon loss, in terms of both the absolute flux and relative stimulation. Consequently, for this scrub–oak ecosystem carbon sequestration was greater in the elevated C_a during this 2‐year study period.     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

A field study of the effects of elevated CO2 on carbon assimilation, stomatal conductance and leaf and branch growth of Pinus taeda trees

A study was conducted in 21-year-old loblolly pine (Pinus taeda L.) trees growing in plantation in north central Georgia, USA. The experiment used branch chambers to impose treatments of ambient, ambient +165 and ambient + 330 μmol mol^?1 CO₂. After one growing season there was no indication of acclimation to elevated CO₂. In August and September, carbon assimilation, measured by two different methods, was twice as high at ambient +330 μmol mol^?1 CO₂ than at ambient. Dark respiration was suppressed by 6% at ambient +165 and by 14% at ambient + 330 μmol mol^?1 CO₂. This suppression was immediate, and not an effect of exposure to elevated CO₂ during growth, since respiration was reduced by the same amount in all treatments when measured at a high CO₂ concentration. Elevated CO₂ increased the growth of foliage and woody tissue. It also increased instantaneous transpiration efficiency, but it had no effect on stomatal conductance. Since the soil at the study site had low to moderate fertility, these results suggest that the growth potential of forests on many sites may be enhanced by global increases in atmospheric CO₂, concentration.     

Acclimation of photosynthesis and respiration to elevated atmospheric CO2 in two Scrub Oaks

Abstract For two species of oak, we determined whether increasing atmospheric CO₂ concentration (C_a) would decrease leaf mitochondrial respiration (R) directly, or indirectly owing to their growth in elevated C_a, or both. In particular, we tested whether acclimatory decreases in leaf‐Rubisco content in elevated C_a would decrease R associated with its maintenance. This hypothesis was tested in summer 2000 on sun and shade leaves of Quercus myrtifolia Willd. and Quercus geminata Small. We also measured R on five occasions between summer 1999 and 2000 on leaves of Q. myrtifolia. The oaks were grown in the field for 4 years, in either current ambient or elevated (current ambient + 350 µmol mol^?1) C_a, in open‐top chambers (OTCs). For Q. myrtifolia, an increase in C_a from 360 to 710 µmol mol^?1 had no direct effect on R at any time during the year. In April 1999, R in young Q. myrtifolia leaves was significantly higher in elevated C_a—the only evidence for an indirect effect of growth in elevated C_a. Leaf R was significantly correlated with leaf nitrogen (N) concentration for the sun and shade leaves of both the species of oak. Acclimation of photosynthesis in elevated C_a significantly reduced maximum RuBP‐saturated carboxylation capacity (V_{c max}) for both the sun and shade leaves of only Q. geminata. However, we estimated that only 11–12% of total leaf N was invested in Rubisco; consequently, acclimation in this plant resulted in a small effect on N and an insignificant effect on R. In this study measurements of respiration and photosynthesis were made on material removed from the field; this procedure had no effect on gas exchange properties. The findings of this study were applicable to R expressed either per unit leaf area or unit dry weight, and did not support the hypothesis that elevated C_a decreases R directly, or indirectly owing to acclimatory decreases in Rubisco content.     

The effect of open-air fumigation with SO2 and O3 on carbohydrate metabolism in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies)

The carbohydrate metabolism of the needles of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) has been examined in trees that were exposed to SO₂, and O₃, in an open-air fumigation experiment located in the Liphook forest in southern England. Two-year-old seedlings were planted in 1985 in seven experimental plots. Five plots received fumigation treatments of SO₂, O₃ or a combination of these gases to give a 2 × 3 factorial design with one additional ambient plot Fumigation with SO₂, occurred from May 1987 to December 1990 and O₃, fumigation occurred from March to December 1988, May to December 1989 and February to December 1990. Five samples of needles for investigation of carbohydrate metabolism were taken between February and July 1989. The concentrations of soluble carbohydrates (including sucrose and hexoses) were greatly reduced in the needles taken from Scots pine growing in the treated plots, and were also reduced, but to a lesser extent, in the needles taken from Norway spruce. Little variation in the concentration of starch in the needles of either species was detected. The activities of the two final enzymes of sucrose synthesis, sucrose phosphate synthase and sucrose 6-phos-phate phosphatase, were greatly reduced in the needles of Scots pine and were also reduced, but to a lesser extent, in the needles of Norway spruce in the fumigated plots. These reductions could be correlated with decreases in rates of photosynthetic CO₂ assimilation determined by independent groups of researchers working on the Liphook site.     

Persistent effects of low concentrations of SO2 and NO2 on photosynthesis in Scots pine (Pinus sylvestris) needles

In an open-field experiment, 50-year-old trees of Scots pine (Pinus sylvestris L.) were fumigated with low concentrations of SO₂ and NO₂ (10–15 nl I^?1) during the growing season in four consecutive years (1988 to 1991). Results from the autumn and early winter of 1991 and 1992 are presented. The maximum photochemical efficiency of photosystem II (PSII), as indicated by the ratio of variable to maximum fluorescence (F_v/F_M) was assessed in current and one-year-old needles from the top and the bottom of the canopy. Furthermore, simultaneous measurements of photosynthetic O₂ evolution and chlorophyll fluorescence were made in current-year needles at 20°C. In general, the F_v/F_M ratio as well as the gross rate of O₂ evolution in needles of fumigated trees was not significantly different from that in needles of control trees during the fumigation period. However, both current and one-year-old needles sampled in November and December 1991 from the top of the canopy of fumigated trees had significantly lower F_v/F_M values than corresponding needles of control trees. Similar differences in F_v/F_M correlated with the treatments were observed in needles from the bottom of the canopy, indicating that the depression of F_v/F_M in needles of fumigated trees was not due to an increased susceptibility to photoinhibition. In 1992, when no fumigation occurred, differences in F_v/F_M between the treatments were not significant during autumn and early winter. The gross rate of O₂ evolution at high irradiances was significantly lower in current-year needles of fumigated trees sampled in November and December 1991 than in those of control trees. Furthermore, a nearly identical linear relationship between the quantum yield of PSII electron transport determined from chlorophyll fluorescence and the quantum yield of O₂ evolution (gross rate of O₂ evolution/PPFD) was found during autumn and early winter. This appeared to be largely a result of changes in the thermal energy dissipation within PSII. The observed differences in photosynthetic characteristics correlated with the different treatments after the fumigation period is suggested to be mainly caused by increased sensitivity of the needles of fumigated trees to low and subfreezing temperatures. However, current-year needles of fumigated trees tended to have a lower N content than those of control trees, which may partly explain the differences in gross photosynthesis between fumigated and control trees.     

Effects of elevated CO2 concentration on photosynthesis, respiration and carbohydrate status of coppice Populus hybrids

To determine how increased atmospheric CO₂ will affect the physiology of coppiced plants, sprouts originating from two hybrid poplar clones ( Populus trichocarpa × P. deltoides - Beaupre and P. deltoides × P. nigra - Robusta) were grown in open-top chambers containing ambient or elevated (ambient + 360 μmol mol⁻¹) CO₂ concentration. The effects of elevated CO₂ concentration on leaf photosynthesis, stomatal conductance, dark respiration, carbohydrate concentration and nitrogen concentration were measured. Furthermore, dark respiration of leaves was partitioned into growth and maintenance components by regressing specific respiration rate vs specific growth rate. Sprouts of both clones exposed to CO₂ enrichment showed no indication of photosynthetic down-regulation. During reciprocal gas exchange measurements, CO₂ enrichment significantly increased photosynthesis of all sprouts by approximately 60% ( P < 0.01) on both an early and late season sampling date, decreased stomatal conductance of all sprouts by 10% ( P < 0.04) on the early sampling date and nonsignificantly decreased dark respiration by an average of 11%. Growth under elevated CO₂ had no consistent effect on foliar sugar concentration but significantly increased foliar starch by 80%. Respiration rate was highly correlated with both specific growth rate and percent nitrogen. Long-term CO₂ enrichment did not significantly affect the maintenance respiration coefficient or the growth respiration coefficient. Carbon dioxide enrichment affected the physiology of the sprouts the same way it affected these plants before they were coppiced.     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

19.

Uptake of SO₂ in shoots of Scots pine: field measurements of net flux of sulphur in relation to stomatal conductance   总被引：2，自引：0，他引：2  

J-E. HÄLLGREN  S. LINDER  A. RICHTER  E. TROENG  L. GRANAT 《Plant, cell & environment》1982,5(1):75-83

Abstract. Net fluxes of gaseous sulphur compounds, water vapour and carbon dioxide to current- and one-year-old shoots of Scots pine were simultaneously measured in the field. The shoots were fumigated in temperature-controlled assimilation chambers with sulphur dioxide at concentrations ranging from 40 to 250 μg m⁻³ (15–95 ppb). The hypothesis that stomatal opening regulates the uptake of sulphur dioxide was tested. The following conclusions were reached concerning the dry deposition of sulphur dioxide to a dry coniferous shoot.
There was a marked diurnal variation in the uptake rate of sulphur dioxide. The net deposition velocity of sulphur was about three times higher during the day than during the night during July-October. Stomatal opening was not the primary factor controlling the dry deposition rate of sulphur dioxide. The net dry deposition rate was significantly smaller than the calculated rate based on stomatal conductances for water vapour.
Part of this deviation could be explained by re-emission of reduced sulphur compound(s) from the needles. The re-emission of sulphur was a light-dependent process and might be of great significance for the sulphur balance calculations.
A variable deposition velocity should be used in atmospheric transport models to account for the diurnal and seasonal variation in the surface resistance of a dry canopy.     

20.

Fine-root respiration in a loblolly pine and sweetgum forest growing in elevated CO₂     

K. George  R. J. Norby  J. G. Hamilton  E. H. DeLucia 《The New phytologist》2003,160(3):511-522

     

Uptake of SO2 in shoots of Scots pine: field measurements of net flux of sulphur in relation to stomatal conductance

Abstract. Net fluxes of gaseous sulphur compounds, water vapour and carbon dioxide to current- and one-year-old shoots of Scots pine were simultaneously measured in the field. The shoots were fumigated in temperature-controlled assimilation chambers with sulphur dioxide at concentrations ranging from 40 to 250 μg m⁻³ (15–95 ppb). The hypothesis that stomatal opening regulates the uptake of sulphur dioxide was tested. The following conclusions were reached concerning the dry deposition of sulphur dioxide to a dry coniferous shoot.
There was a marked diurnal variation in the uptake rate of sulphur dioxide. The net deposition velocity of sulphur was about three times higher during the day than during the night during July-October. Stomatal opening was not the primary factor controlling the dry deposition rate of sulphur dioxide. The net dry deposition rate was significantly smaller than the calculated rate based on stomatal conductances for water vapour.
Part of this deviation could be explained by re-emission of reduced sulphur compound(s) from the needles. The re-emission of sulphur was a light-dependent process and might be of great significance for the sulphur balance calculations.
A variable deposition velocity should be used in atmospheric transport models to account for the diurnal and seasonal variation in the surface resistance of a dry canopy.