Ecology of SO2 resistance: I. Effects of fumigations on gas exchange of deciduous and evergreen shrubs

Summary A unique gas exchange system is described in which photosynthesis, transpiration, and stomatal conductance can be measured on leaves during SO₂ fumigations. SO₂ concentrations can be continuously monitored and manipulated between 0 and 2.0 ppm. Rates of total SO₂ uptake and SO₂ absorption through stomates of a fumigated leaf can also be determined.Using this system we compared the effects of SO₂ on the gas exchange rates of two shrub species that co-occur in the Califormian chaparral. Diplacus aurantiacus, a deciduous shrub, was more sensitive to SO₂ fumigation than Heteromeles arbutifolia, an evergreen shrub. The differences in photosynthetic sensitivity could be attributed, in large part, to differential SO₂ absorption rates.     

Search for polyprenols in leaves of evergreen and deciduous Ericaceae plants.

Various species and cultivars of Ericaceae family were checked for the presence of long-chain polyprenols in their leaves. In the genus Rhododendron no polyprenols were found in the ever-green species, while they were present in the deciduous type. The polyprenols were of chain-length of 14-20 isoprene residues and they occurred in the form of acetic acid esters. The polyprenol accumulation is discussed with respect to senescence of leaves.     

Energy content,storage substances,and construction and maintenance costs of Mediterranean deciduous leaves

Summary At monthly intervals water content, crude fibre, total and protein nitrogen, sugars, starch, total lipids, ash content and calorific total energy were measured throughout the lifespan of the leaves of the deciduous mediterranean shrubs Pistacia terebinthus L. and Cotinus coggygria Scop. From these data the construction costs and maintenance costs, as well as the construction costs of non-storage compounds and energy expenditure values were calculated. The latter values were also calculated for the evergreen stemmed shrub Ephedra distachya for reasons of comparison with an evergreen mediterranean species. The water status in the deciduous leaves is stable for a long time during the drought period until the beginning of senescence in August/September. In Cotinus an early and considerable increase of storage compounds is found, whilst in Pistacia terebinthus the accumulation is more uniform until August. The N-content is rather low compared with other deciduous leaves, the calorific energy is in the lower range of the values reported for similar species. The construction costs of the leaves of both deciduous species are significantly lower than those calculated by Williams et al. (1987) for two drought deciduous chaparral species but are in agreement with the data reported by other authors on deciduous leaves. Contrary to the findings of Williams et al. they are lower than those of evergreen species; this is also true when the construction cost of the non-storage compounds alone is considered. The values found for Ephedra are similar to the maintenance costs do not show a significant variation in the deciduous leaves. They are higher than those known for evergreen leaves but somewhat lower than the values calculated for deciduous chaparral leaves.     

Ultrasound acoustic emissions from dehydrating leaves of deciduous and evergreen trees

While drying, detached leaves produced ultrasound acoustic emissions (UAE) comparable to emissions from stem and twig wood. Experiments on Ilex aquifolium L. showed that the main source of these signals was cavitation in the veins, to which conduits and fibres probably both contributed. Regions of the leaf blade with abundant mesophyll and only small veins emitted few signals. More signals were counted on the adaxial side of the midrib than on the abaxial one and on the proximal third than on the distal one, in accordance with the anatomical structure. Sound attenuation was pronounced. Eight species were compared with respect to cavitation behaviour, field water relations and pressure–volume curves, and the data showed differences in cumulative number of events and resistance of leaves to cavitation. Data were generally in good agreement with anatomical structure and habitat preferences. The number of signals per conduit counted on cross-sections was in some leaves much higher than unity, which suggests short xylem elements or an acoustic activity of cells other than conduits. There was no correlation between cavitation threshold or cumulative number of signals and the degree of sclerophylly; unexpectedly, there was a correlation between the cumulative number of signals at a water potential of -1.3 MPa and the bulk modulus of elasticity.     

On the difference in the net ecosystem exchange of CO2 between deciduous and evergreen forests in the southeastern United States

The southeastern United States is experiencing a rapid regional increase in the ratio of pine to deciduous forest ecosystems at the same time it is experiencing changes in climate. This study is focused on exploring how these shifts will affect the carbon sink capacity of southeastern US forests, which we show here are among the strongest carbon sinks in the continental United States. Using eight‐year‐long eddy covariance records collected above a hardwood deciduous forest (HW) and a pine plantation (PP) co‐located in North Carolina, USA, we show that the net ecosystem exchange of CO₂ (NEE) was more variable in PP, contributing to variability in the difference in NEE between the two sites (ΔNEE) at a range of timescales, including the interannual timescale. Because the variability in evapotranspiration (ET) was nearly identical across the two sites over a range of timescales, the factors that determined the variability in ΔNEE were dominated by those that tend to decouple NEE from ET. One such factor was water use efficiency, which changed dramatically in response to drought and also tended to increase monotonically in nondrought years (P < 0.001 in PP). Factors that vary over seasonal timescales were strong determinants of the NEE in the HW site; however, seasonality was less important in the PP site, where significant amounts of carbon were assimilated outside of the active season, representing an important advantage of evergreen trees in warm, temperate climates. Additional variability in the fluxes at long‐time scales may be attributable to slowly evolving factors, including canopy structure and increases in dormant season air temperature. Taken together, study results suggest that the carbon sink in the southeastern United States may become more variable in the future, owing to a predicted increase in drought frequency and an increase in the fractional cover of southern pines.     

Root respiration and associated costs in deciduous and evergreen species of Quercus

Root construction and maintenance costs were estimated in four evergreen and three deciduous Quercus species that are typical in the landscape of southern Spain. The cost quantification was based on analysis of the growth–­respiration ratio. Values observed for both construction cost (ranging from 1·17 to 1·29 g glucose g^?1 dry weight) and maintenance cost (ranging from 6·22 to 11·71 mg glucose g^?1 dry weight d^?1) were generally lower than those reported in other studies. The results showed non‐significant differences between deciduous and evergreen species. The lack of significant differences between species appeared to be due to the homogeneity of growth conditions. Hydroponic culture, with unrestricted nutrient and water supply, would lead to low tissue carbon content and low respiration rates, leading to the low costs observed. Furthermore, the fact that root organs are clearly importers of organic molecules inevitably entails some underestimation of the respiration associated with growth and, to a lesser extent, with maintenance respiration. This leads in turn to underestimation of the corresponding construction and maintenance costs. All this raises doubts as to the suitability of this method for studying root systems.     

Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species

Photosynthetic nitrogen use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) is one of the most important factors for the interspecific variation in photosynthetic capacity. PNUE was analysed in two evergreen and two deciduous species of the genus Quercus. PNUE was lower in evergreen than in deciduous species, which was primarily ascribed to a smaller fraction of nitrogen allocated to the photosynthetic apparatus in evergreen species. Leaf nitrogen was further analysed into proteins in the water‐soluble, the detergent‐soluble, and the detergent‐insoluble fractions. It was assumed that the detergent‐insoluble protein represented the cell wall proteins. The fraction of nitrogen allocated to the detergent‐insoluble protein was greater in evergreen than in deciduous leaves. Thus the smaller allocation of nitrogen to the photosynthetic apparatus in evergreen species was associated with the greater allocation to cell walls. Across species, the fraction of nitrogen in detergent‐insoluble proteins was positively correlated with leaf mass per area, whereas that in the photosynthetic proteins was negatively correlated. There may be a trade‐off in nitrogen partitioning between components pertaining to productivity (photosynthetic proteins) and those pertaining to persistence (structural proteins). This trade‐off may result in the convergence of leaf traits, where species with a longer leaf life‐span have a greater leaf mass per area, lower photosynthetic capacity, and lower PNUE regardless of life form, phyllogeny, and biome.     

Contrasting acclimation responses to elevated CO2 and warming between an evergreen and a deciduous boreal conifer

Rising atmospheric carbon dioxide (CO₂) concentrations may warm northern latitudes up to 8°C by the end of the century. Boreal forests play a large role in the global carbon cycle, and the responses of northern trees to climate change will thus impact the trajectory of future CO₂ increases. We grew two North American boreal tree species at a range of future climate conditions to assess how growth and carbon fluxes were altered by high CO₂ and warming. Black spruce (Picea mariana, an evergreen conifer) and tamarack (Larix laricina, a deciduous conifer) were grown under ambient (407 ppm) or elevated CO₂ (750 ppm) and either ambient temperatures, a 4°C warming, or an 8°C warming. In both species, the thermal optimum of net photosynthesis (T_optA) increased and maximum photosynthetic rates declined in warm‐grown seedlings, but the strength of these changes varied between species. Photosynthetic capacity (maximum rates of Rubisco carboxylation, V_cmax, and of electron transport, J_max) was reduced in warm‐grown seedlings, correlating with reductions in leaf N and chlorophyll concentrations. Warming increased the activation energy for V_cmax and J_max (E_aV and E_aJ, respectively) and the thermal optimum for J_max. In both species, the T_optA was positively correlated with both E_aV and E_aJ, but negatively correlated with the ratio of J_max/V_cmax. Respiration acclimated to elevated temperatures, but there were no treatment effects on the Q₁₀ of respiration (the increase in respiration for a 10°C increase in leaf temperature). A warming of 4°C increased biomass in tamarack, while warming reduced biomass in spruce. We show that climate change is likely to negatively affect photosynthesis and growth in black spruce more than in tamarack, and that parameters used to model photosynthesis in dynamic global vegetation models (E_aV and E_aJ) show no response to elevated CO₂.     

CO2 gas exchange and phosphoenolpyruvate carboxylase activity in leaves of Zea mays L.

Important gas exchange characteristics of C4 plants depend on the properties of phophoenolpyruvate carboxylase (PEPC), the enzyme catalysing the primary fixation of CO2 during C4 photosynthesis. In this study, the relationship between intracellular resistance for CO2 fixation (r_i) at high photosynthetically active photon flux densities (PPFD) and maximum PEPC activity in vitro (V_pm) was examined in leaves of Zea mays L. The analysis allowed the estimation of the Michaelis constant K_p of the enzyme for CO2 (or the equivalent number for bicarbonate) in vivo. At low PPFD (below 100 mol m-2 s-1) the initial slopes of the curves describing net CO2 uptake rate A as a function of intercellular CO2 concentration c_i increased with increasing PPFD. The increase (i. e. a decrease in r_i) was interpreted as due to a reversible activation of PEPC by light. Including this assumption into a model of C4 photosynthesis enabled us to reproduce A(c_i) response curves measured at low levels of PPFD. Fitting the model to experimental data resulted in values for K_I, the PPFD at which PEPC reaches half of its full activation, of about 200 mol m-2 s-1. Similar results were derived from the dependence of r_i on PPFD. The analysis of the relationships between r_i and V_pm and between r_i and PPFD, as well as fitting of the model to gas exchange data all gave rise to estimates for the resistance for CO2 transfer within mesophyll cells that are comparable with those known from C3 plants.     

Construction costs and mesostructure of leaves in hydrophytes

Construction costs (CC) and parameters of leaf structure (specific leaf weight, dry matter content, volume of photosynthesizing cells, and the number of cells per leaf area unit) were determined for 19 species of aquatic higher plants. The CC of 1 g dry matter varied from 0.98 g glucose in Lemna gibba L. to 1.48 g glucose in Nuphar pumila (Timm) DC. and Potamogeton natans L. The CC of leaf area unit varied to a greater extent than the CC of 1 g dry wt (from 10 to 97 g glucose/m²) and depended on the type of mesophyll structure. In leaves of hydrophytes with dorsoventral mesophyll structure, the CC of 1 m² leaf area was 3–9 times larger than in leaves with homogeneous structure. Variations in CC of 1 m² leaf area in hydrophytes were affected insignificantly (by 2% only) by variations of CC per 1 g dry wt and were mainly determined (by 82%) by changes in specific leaf weight. Two-factor analysis of variance has shown that the CC of 1 g dry wt in hydrophytes depended on the attachment of plants to the sediment: the CC was 1.2 times larger in rooted hydrophytes than in free floating plants. The second factor (the extent of submergence) potentiated the effect of rooting on CC. Reliable differences were found between the leaf CC for hydrophytes belonging to four groups distinguished by the extent of their contact with water and sediment. In a group series: rooted hydrophytes with floating leaves → submerged rooted hydrophytes → free floating submerged hydrophytes → free floating surface inhabiting hydrophytes, the CC of 1 g dry wt decreased by 1.3 times. Path analysis has shown that this trend was due to the increase in photosynthesizing cell volume and to reduction in number of cells per leaf area unit, which caused the decrease in dry matter content. The decrease in the content of leaf dry matter was accompanied by changes in its chemical composition: the content of carbon and nitrogen decreased. This led to a consistent decrease in leaf CC expressed per 1 g dry wt upon the increase in extent of plant hydrophilicity.     

Deriving a new phenological indicator of interannual net carbon exchange in contrasting boreal deciduous and evergreen forests

Phenology is an important variable affecting the annual net ecosystem production (NEP) of terrestrial ecosystems. A new phenological indicator was proposed based on the ratio of respiration season length and growing season length (respiration–growth length ratio, RGR). Validation of this new phenological indicator was conducted using continuous flux measurements at contrasting boreal deciduous and evergreen forests in Canada. Analyses based on yearly anomalies of both annual NEP and phenological indicators indicated that the RGR can explain more proportion of interannual NEP variability compared to existing phenological metrics, including the carbon uptake period and the autumn lag. A multivariate regression model was used to predict the respiration–growth length ratio anomaly using anomalies of spring air temperature, autumn radiation and soil water content (SWC), which serves as a prerequisite for this indicator being scaled up for regional applications where flux data were unavailable. By normalization growing season length, interannual NEP showed comparable sensitivity to RGR variations of different plant functional types, which is a great advantage over other phenological indicators. The high potential of RGR in explaining interannual NEP variability may highlight the importance of respiration process in controlling annual NEP, which has probably been overlooked or underestimated in existing phenological studies. The comparable sensitivity of RGR to annual NEP observed at different plant functional types would favor its application in tracking interannual variability of NEP regionally and complementary to existing indices to promote our understanding of carbon sequestration with future climate change.     

Comparative vessel anatomy of arctic deciduous and evergreen dicots

Arctic tundra plant species exhibit striking variation in leaf character and growth form. Both are likely related to differences in vessel anatomy, and all may affect responses to climate changes in the Arctic. To investigate the relationships among leaf character, growth form, vessel anatomy, and susceptibility to freeze-thaw-induced xylem cavitation, xylem vessel characteristics were compared among six deciduous and six evergreen arctic dicot species of erect and prostrate growth forms. We hypothesized that deciduous and erect species would have larger and longer vessels than evergreen and cushion/mat-forming species. Vessel lengths, diameters, and densities were measured for each species. Theoretical vessel flow rates were calculated using Poiseuille's law for ideal capillaries. Flow rates were used to determine the susceptibility of vessels to cavitation induced by freeze-thaw events that may become more frequent with global warming. Vessel diameters were larger in deciduous species compared to evergreens, and in shrubs/trees vs. cushion/mat-forming plants. Vessel length distributions, however, did not differ for growth form or leaf character. Vessel density was greater in cushion/mat-forming species than in shrub/tree species. Deciduous plants showed a greater contribution to total conductivity by relatively larger vessels than evergreens. One of the deciduous species, Vaccinium uliginosum, is predicted to be susceptible to freeze-thaw-induced cavitation. These results have important implications for future arctic species composition and plant community structure.     

Underwater photosynthesis and respiration in leaves of submerged wetland plants: gas films improve CO2 and O2 exchange.

Many wetland plants have gas films on submerged leaf surfaces. We tested the hypotheses that leaf gas films enhance CO(2) uptake for net photosynthesis (P(N)) during light periods, and enhance O(2) uptake for respiration during dark periods. Leaves of four wetland species that form gas films, and two species that do not, were used. Gas films were also experimentally removed by brushing with 0.05% (v/v) Triton X. Net O(2) production in light, or O(2) consumption in darkness, was measured at various CO(2) and O(2) concentrations. When gas films were removed, O(2) uptake in darkness was already diffusion-limited at 20.6 kPa (critical O(2) pressure for respiration, COP(R)>/= 284 mmol O(2) m(-3)), whereas for some leaves with gas films, O(2) uptake declined only at approx. 4 kPa (COP(R) 54 mmol O(2) m(-3)). Gas films also improved CO(2) uptake so that, during light periods, underwater P(N) was enhanced up to sixfold. Gas films on submerged leaves enable continued gas exchange via stomata and thus bypassing of cuticle resistance, enhancing exchange of O(2) and CO(2) with the surrounding water, and therefore underwater P(N) and respiration.     

Effects of CO2 Enrichment on Four Poplar Clones. I. Growth and Leaf Anatomy

The poplar clones Columbia River, Beaupre, Robusta and Raspaljehave been investigated under the present (350 µmol mol^–1)and double the present (700 µmol mol^–1) atmosphericCO₂ concentration. Cuttings were planted in pots and were grownin open-top chambers inside a glasshouse for 92 d. The number of leaves, total length of stem, total leaf area,overall growth rate, total leaf, stem and root d. wt respondedpositively to increased CO₂ but the leaf size and biomass allocationshowed no change with CO₂ enrichment. Beaupre and Robusta showeda larger growth response than either Columbia River or Raspalje. The effects of CO₂ enrichment were restricted to the early phaseof growth at the beginning of the growth season. Leaf cell numbers in all the clones were not affected by CO₂enrichment. Leaf thickness was affected; this was mainly theresult of larger mesophyll cells and more extensive intercellularspaces. Poplar clones, CO₂ enrichment, growth, leaf anatomy, leaf cell number     

Net CO2 exchange rates and diffusion resistance of leaves from chicory selections during vegetative growth

Net CO₂ exchange rates, stomatal and internal resistances for CO₂-transport were followed on fully expanded Witloof chicory leaves (Cichorium intybus L. cv. Foliosum) for several months during vegetative growth. Maximum net CO₂ exchange rate (P_max) stayed high with a sudden drastic drop at the end of the growing season largely due to an increase in internal diffusion resistance. During an analogous growth period the H₂O vapour diffusion resistances of leaves for four selections were measured. The adaxial stomatal resistance was always higher than the abaxial one. Stomatal densities calculated for those selections showed higher values at the abaxial leaf side.     

Low stomatal and internal conductance to CO2 versus Rubisco deactivation as determinants of the photosynthetic decline of ageing evergreen leaves

A novel A-Ci curve (net CO2 assimilation rate of a leaf -An- as a function of its intercellular CO2 concentration -Ci) analysis method (Plant, Cell & Environment 27, 137-153, 2004) was used to estimate the CO2 transfer conductance (gi) and the maximal carboxylation (Vcmax) and electron transport (Jmax) potentials of ageing, non-senescing Pseudotsuga menziesii leaves in relation to their nitrogen (N) content and protein and pigment composition. Both gi and the stomatal conductance (gsc) of leaves were closely coupled to Vcmax, Jmax and An with all variables decreasing with increasing leaf age. Consequently, both Ci and Cc (chloroplastic CO2 concentration) remained largely conserved through successive growing seasons. The N content of leaves, as well as the amount of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and other sodium dodecyl sulfate-soluble proteins, increased during the first three growing seasons, then stabilized or decreased only slightly afterwards. Thus, the age-related photosynthetic nitrogen use efficiency (PNUE) decline of leaves was not a consequence of decreased allocation of N towards Rubisco and other proteins involved in bioenergetics and light harvesting. Rather, loss of photosynthetic capacity was the result of the decreased activation state of Rubisco and proportional down-regulation of electron transport towards the photosynthetic carbon reduction (PCR) and photorespiratory (PCO) cycles in response to a reduction of CO2 supply to the chloroplasts' stroma. This study emphasizes the regulatory potential and homeostaticity of Cc- rather than photosynthetic metabolites or Ci- in relation to the commonly observed correlation between photosynthesis and gsc.     

An analysis of the temperature response curves of CO2 exchange in the leaves of two temperate and one tropical grass species

Summary The temperature dependence of carbon dioxide exchange in two temperate grass species, Lolium perenne (S.23) and Lolium multiflorum (S.22), and one tropical grass species, Cenchrus ciliaris, have been examined. It has been shown that the decline in the net CO₂ exchange rate above the optimum temperature is caused, in the case of C. ciliaris, by denaturation, and, in the case of L. perenne and L. multiforum by the greater response to temperature of respiration compared with photosynthesis.Activation energies have been obtained for both photosynthetic and respiratory systems. For both systems the values fall into discrete groups, and it is postulated that each group may be characteristic of a particular rate-limiting process. The distribution of values changes with changes in experimental conditions, there being marked differences in the distribution of values obtained for light-limited and CO₂-limited photosynthesis.     

Differences in construction costs and chemical composition between deciduous and evergreen woody species are small as compared to differences among families

We tested to what extent differences in construction costs (CC) and chemical composition of woody species are attributed to leaf habit. Eight evergreen and eight deciduous species belonging to six families were selected to form eight phylogenetic independent contrasts (PICs). The plants were grown from seed in a glasshouse. Differences in leaf, stem and root CC between evergreen and deciduous species were minor, the proportion of variance explained by leaf habit generally being less than 6%. Surprisingly, differences in leaf chemical composition between deciduous and evergreen species were small as well. Variation in CC and chemical composition among families was substantial, the factor 'family' explaining 50-85% of variance. We therefore conclude that in this case, phylogeny is a more important factor than functional group. Leaves of the fast-growing species in this experiment showed high levels of minerals, organic acids, proteins and lipids, whereas leaves of the slow-growing species had higher concentrations of soluble phenolics, lignin as well as higher carbon/nitrogen (C/N) ratio. These relationships suggest a trade-off between growth and defence. In contrast, CC of leaves, stems, roots or whole plants showed no or only a weak correlation with relative growth rate (RGR). The C/N ratio of the leaves is an easily measured parameter that correlated strongly in a negative way with the RGR of the plants and reflected better the balance between investment in structure and physiological functioning than CC.     

叶片和群落尺度净光合速率关系的探讨

叶片净光合速率(P_n)是研究光合作用机理的基本尺度; 而群落净光合速率(P_c)是研究群落光合能力及其与外部环境因子间关系的更好尺度, 特别是区域乃至全球尺度碳循环的研究, 需要将叶片尺度的生理生态模型扩展到冠层尺度。理论上, 群落内所有叶片的累积P_n与实测群落净气体交换速率(NCE)是相等的, 但在野外实际观测中, 两者之间的相互关系目前尚未见报道。该文选取敖汉苜蓿(Medicago sativa ‘Aohan’)人工草地, 采用美国LI-COR公司生产的便携式光合测定系统LI-6400测定P_n, 结合叶面积指数等参数推算P_c, 利用LI-8100连接同化箱测定生态系统净气体交换速率(NEE), 加上土壤呼吸速率, 得到NCE。结果表明: P_c为3.52 μmol CO₂·m^-2·s^-1, 与实测NCE (3.56 μmol CO₂·m^-2·s^-1)基本相等。这表明: 可利用P_n, 结合叶面积指数、群落叶片数目、健康叶片比例和群落可接收有效光照的平均比例等4个关键参数, 准确地换算P_c。然而, 利用同化箱式法测定群落呼吸速率时, 不可避免地会包含土壤呼吸, 所以在观测NCE时, 需要同时测定土壤呼吸。此外, 在冠层模型中, 群落垂直结构和光量子的非线性响应不可忽视。