Influence of shoot structure on light interception and photosynthesis in conifers

The influence of shoot structure on net photosynthesis was evaluated under field conditions for the central Rocky Mountain (United States) conifers Picea engelmannii (Parry ex Engelm.), Abies lasiocarpa ([Hook] Nutt.), and Pinus contorta (Engelm.). In all species, the greater number of needles per unit stem length on sun shoots correlated with a smaller silhouette leaf area to total leaf area ratio (STAR). Decreased STAR was due primarily to greater needle inclination toward the vertical, plus some needle mutual shading. However, photosynthesis expressed on a total leaf area basis did not decrease in sun shoots (lower STAR) but remained nearly constant at approximately 3 micromoles per square meter per second over a wide range of STAR (0.1 to 0.3). Relatively low light saturation levels of 200 to 1400 microeinsteins per square meter per second and diffuse light to 350 microeinsteins per meter per second maintained photosynthetic flux densities in inclined and/or shaded needles at levels comparable to those in unshaded needles oriented perpendicular to the solar beam. As a result, net CO₂ uptake per unit stem length increased as much as 2-fold in sun shoots (low STAR) in direct proportion to increasing needle density.     

Influence of krummholz mat microclimate on needle physiology and survival

Summary Microclimate and photosynthesis of krummholz mat growth forms of Picea engelmanii (Parry) and Abies lasiocarpa [Hook.] Nutt. were investigated to determine structural features which may aid survival in alpine environments. The structure of krummholz mats was described in terms of the vertical distribution of leaf area index and leaf area density, which exceeded 50 m^-1 (based on total leaf surface area) near the canopy surface and approached zero below 30 cm from the surface in both species. Photosynthetic photon flux density (PPFD, 0.4–0.7 m wavelengths) and wind decreased by an average of 6 and 50-fold, respectively, between 1 m above and 10 cm below mat surfaces in both species. Needle temperatures on a P. engelmannii krummholz mat during July averaged about 2°C above air temperature during the day, with a maximum overtemperature of greater than 20°C above T _air during one sunlit period. At night, needle temperatures averaged 3–4°C below T _air.Net photosynthesis in year-old P. engelmannii shoots reached a maximum at 15–20°C during July and August. Surface shoots were light saturated at near 1200 moles m^-2s^-1 PPFD, and had higher photosynthetic rates than subsurface, predominantly shaded shoots above 800 moles m^-2s^-1. Shade shoots had higher photosynthetic rates when PPFD was below 600 moles m^-2s^-1, and at 250 moles m^-2s^-1 shade shoots maintained about 50% of the net photosynthetic rate of sun shoots at light saturation. Shade shoots appeared capable of benefitting photosynthetically from elevated temperatures within krummholz mats despite relatively low light levels. Especially rapid photosynthesis may occur when canopy needles are illuminated by sunflecks and needle temperatures rise by 10° C or more.Snow cover appears crucial for the survival of needles during winter. Snow accumulated within krummholz needle canopies before the sub-canopy zone of unfoliated branches became filled. The concentrated needle growth in the krummholz canopy captured snow in early autumn without support from ground-level snowpack. Early snow cover in both species prevented cuticle abrasion and resulted in high winter needle water contents and viabilities for subsurface compared to surface needles which became abraded, severely dehydrated, and had high mortality between December and February, especially on windward sides of shoots.Extremely high concentrations of needles within krummholz mat canopies created an aerodynamic structure which elevated needle temperatures to more optimal photosynthetic levels in summer and resulted in more efficient snow accumulation in winter. These factors appear crucial for winter needle survival. Thus, krummholz mats appear to be an important adaptation in growth form which provides survival benefits in both summer and winter.     

Seasonal changes in temperature response of photosynthesis and its contribution to annual carbon gain in Daphniphyllum humile,an evergreen understorey shrub

We evaluated seasonal variation in photosynthetic temperature dependence and its contribution to annual carbon gain in an evergreen understorey shrub, Daphniphyllum humile Maxim, growing at the forest border and in the understorey of a deciduous forest. Plants at both sites exhibited similar optimal temperatures for photosynthesis (T_opt). The activation energy for ribulose‐1,5‐bisphosphate (RuBP) carboxylation (H_aV) at both sites tended to be higher in summer than in spring or autumn, suggesting that H_aV may be the controlling factor in the T_opt shift in D. humile. In contrast to the seasonal changes in T_opt, the maximum photosynthetic rate at the optimal temperature (P_opt) differed between the two sites: it was lower in autumn than in summer at the forest border, but was the same in summer and autumn in the understorey. In the understorey plants, nitrogen content (N_area) increased in autumn, but this was not the case for forest border plants. In addition, Rubisco content increased significantly in autumn in the understorey leaves but decreased distinctly in forest border leaves. Increased N_area and Rubisco in understorey leaves resulted in increased in photosynthesis in autumn. Annual carbon gain was 30.8 mol·m^?2 in forest border leaves and 5.8 mol·m^?2 in understorey leaves. Carbon gain in understorey leaves during the short period after overstorey leaf fall and before snow accumulation was approximately 49% of annual carbon gain. Furthermore, autumn carbon gain calculated using activation energy of summer with autumn photosynthetic parameters underestimated the autumn carbon gain by as much as 31%. In conclusion, photosynthetic temperature acclimation may be a key factor in increasing annual carbon gain in understorey D. humile.     

Gas exchange and canopy structure of 9-year-old loblolly pine,pitch pine and pitch x loblolly hybrids

Summary Seasonal gas exchange and canopy structure were compared among 9-year-old loblolly pine (Pinus taeda L.), pitch pine (Pinus rigida Mill.), and pitch x loblolly hybrids (Pinus rigida x taeda) growing in an F2 plantation located in Critz, Va., USA. Leaf net photosynthesis, conductance, internal CO₂ concentration (c_i), water use efficiency (WUE; photosynthesis/conductance), dark respiration and the ratio of net photosynthesis/respiration did not vary among or within the three taxa. Significant differences in volume production, crown length, total crown leaf surface area and the silhouette area of shade shoots among the taxa were observed. The loblolly-South Carolina source had greater volume and crown surface area than the pitch pine, and the hybrid taxa were intermediate between the two. Although the silhouette area ratio of shade foliage varied among taxa, it was not related to volume. A strong relationship between total leaf surface area and volume was observed. Leaf conductance, c_i, WUE and leaf water potential were the physiological parameters significantly and positively correlated with volume. This study suggests that the amount of needle surface in the canopy is more important in early stand volume growth than the leaf carbon exchange rate and the degree of needle self-shading in the lower canopy.     

Acclimation of photosynthesis,respiration and ecosystem carbon flux of a wetland on Chesapeake Bay,Maryland to elevated atmospheric CO2 concentration

Acclimation of photosynthesis and respiration in shoots and ecosystem carbon dioxide fluxes to rising atmospheric carbon dioxide concentration (C _a ) was studied in a brackish wetland. Open top chambers were used to create test atmospheres of normal ambient and elevated C _a (=normal ambient + 34 Pa CO₂) over mono-specific stands of the C₃ sedge Scirpus olneyi, the dominant C₃ species in the wetland ecosystem, throughout each growing season since April of 1987. Acclimation of photosynthesis and respiration were evaluated by measurements of gas exchange in excised shoots. The impact of elevated C _a on the accumulation of carbon in the ecosystem was determined by ecosystem gas exchange measurements made using the open top chamber as a cuvette.Elevated C _a increased carbohydrate and reduced Rubisco and soluble protein concentrations as well as photosynthetic capacity(A) and dark respiration (R _d ; dry weight basis) in excised shoots and canopies (leaf area area basis) of Scirpus olneyi. Nevertheless, the rate of photosynthesis was stimulated 53% in shoots and 30% in canopies growing in elevated C _a compared to normal ambient concentration. Elevated C _a inhibited R _d measured in excised shoots (–19 to –40%) and in seasonally integrated ecosystem respiration (R _e ; –36 to –57%). Growth of shoots in elevated C _a was stimulated 14–21%, but this effect was not statistically significant at peak standing biomass in midseason. Although the effect of elevated C _a on growth of shoots was relatively small, the combined effect of increased number of shoots and stimulation of photosynthesis produced a 30% stimulation in seasonally integrated gross primary production (GPP). The stimulation of photosynthesis and inhibition of respiration by elevated C _a increased net ecosystem production (NEP=GPP–R _e ) 59% in 1993 and 50% in 1994. While this study consistently showed that elevated C _a produced a significant increase in NEP, we have not identified a correspondingly large pool of carbon below ground.     

Acclimation of photosynthetic temperature optima of temperate and boreal tree species in response to experimental forest warming

Rising temperatures caused by climate change could negatively alter plant ecosystems if temperatures exceed optimal temperatures for carbon gain. Such changes may threaten temperature‐sensitive species, causing local extinctions and range migrations. This study examined the optimal temperature of net photosynthesis (T_opt) of two boreal and four temperate deciduous tree species grown in the field in northern Minnesota, United States under two contrasting temperature regimes. We hypothesized that T_opt would be higher in temperate than co‐occurring boreal species, with temperate species exhibiting greater plasticity in T_opt, resulting in better acclimation to elevated temperatures. The chamberless experiment, located at two sites in both open and understory conditions, continuously warmed plants and soils during three growing seasons. Results show a modest, but significant shift in T_opt of 1.1 ± 0.21 °C on average for plants subjected to a mean 2.9 ± 0.01 °C warming during midday hours in summer, and shifts with warming were unrelated to species native ranges. The 1.1 °C shift in T_opt with 2.9 °C warming might be interpreted as suggesting limited capacity to shift temperature response functions to better match changes in temperature. However, T_opt of warmed plants was as well‐matched with prior midday temperatures as T_opt of plants in the ambient treatment, and T_opt in both treatments was at a level where realized photosynthesis was within 90–95% of maximum. These results suggest that seedlings of all species were close to optimizing photosynthetic temperature responses, and equally so in both temperature treatments. Our study suggests that temperate and boreal species have considerable capacity to match their photosynthetic temperature response functions to prevailing growing season temperatures that occur today and to those that will likely occur in the coming decades under climate change.     

Mathematical models of the photosynthetic response of tree stands to rising CO2 concentrations and temperatures

Two published models of canopy photosynthesis, MAESTRO and BIOMASS, are simulated to examine the response of tree stands to increasing ambient concentrations of carbon dioxide (C_a) and temperatures. The models employ the same equations to described leaf gas exchange, but differ considerably in the level of detail employed to represent canopy structure and radiation environment. Daily rates of canopy photosynthesis simulated by the two models agree to within 10% across a range of CO₂ concentrations and temperatures. A doubling of C_a leads to modest increases of simulated daily canopy photosynthesis at low temperatures (10% increase at 10°C), but larger increases at higher temperatures (60% increase at 30°C). The temperature and CO₂ dependencies of canopy photosynthesis are interpreted in terms of simulated contributions by quantum-saturated and non-saturated foliage. Simulations are presented for periods ranging from a diurnal cycle to several years. Annual canopy photosynthesis simulated by BIOMASS for trees experiencing no water stress is linearly related to simulated annual absorbed photosynthetically active radiation, with light utilization coefficients for carbon of ?= 1.66 and 2.07g MJ^?1 derived for C_a of 350 and 700 μmol mol^?1, respectively.     

Modification of the response of photosynthetic productivity to rising temperature by atmospheric CO2 concentrations: Has its importance been underestimated?

Abstract. Climate change will include correlated increases in temperature and atmospheric CO₂ concentration (C_a). Rising temperatures will increase the ratio of photorespiratory loss of carbon to photosynthetic gain, whilst rising C_a will have an opposing effect. The mechanism of these effects at the level of carboxylation in C₃ photosynthesis are quantitatively well understood and provide a basis for models of the response of leaf and canopy carbon exchange to climate change. The principles of such a model are referred to here and used to quantitatively examine the implications of concurrent increase in temperature and C_a. Simulations of leaf photosynthesis show the increase, with elevation of C_a from 350 to 650 μmol mol^-1, in light saturated rates of CO₂ uptake (A_sat) and maximum quantum yields (φ) to rise with temperature. An increase in C_a from 350 to 650 μmol mol^-1 can increase A_sat by 20% at 10°C and by 105% at 35°C, and can raise the temperature optimum of A_sat by 5°C. This pattern of change agrees closely with experimental data. At the canopy level, simulations also suggest a strong interaction of increased temperature and CO₂ concentration. Predictions are compared with the findings of long-term field studies. The principles used here suggest that elevated C_a will alter both the magnitude of the response of leaf and canopy carbon gain to rising temperature, and sometimes, the direction of response. Findings question the value of models for predicting plant production in response to climate change which ignore the direct effects of rising C_a and the modifications that rising C_a imposes on the temperature response of net CO₂ exchange.     

Carbon uptake and respiration in above-ground parts of a Larix decidua × leptolepis tree

Summary Shade needles of hybrid larch (Larix decidua × leptolepis) had the same rates of photosynthesis as sun needles per dry weight and nitrogen, and a similar leaf conductance under conditions of light saturation at ambient CO₂ (A_max). However, on an area basis, A_max and specific leaf weight were lower in shade than in sun needles. Stomata of sun needles limited CO₂ uptake at light saturation by about 20%, but under natural conditions of light in the shade crown, shade needles operated in a range of saturating internal CO₂ without stomatal limitation of CO₂ uptake. In both needle types, stomata responded similarly to changes in light, but shade needles were more sensitive to changes in vapor pressure deficit than sun needles. Despite a high photosynthetic capacity, the ambient light conditions reduced the mean daily (in summer) and annual carbon gain of shade needles to less than 50% of that in sun needles. In sun needles, the transpiration per carbon gain was about 220 mol mol^–1 on an annual basis. The carbon budget of branches was determined from the photosynthetic rate, the needle biomass and respiration, the latter of which was (per growth and on a carbon basis) 1.6 mol mol^–1 year^–1 in branch and stem wood. In shade branches carbon gains exceeded carbon costs (growth + respiration) by only a factor of 1.6 compared with 3.5 in sun branches. The carbon balance of sun branches was 5 times higher per needle biomass of a branch or 9 times higher on a branch length basis than shade branches. The shade foliage (including the shaded near-stem sun foliage) only contributed approximately 23% to the total annual carbon gain of the tree.     

Plant competition for light analyzed with a multispecies canopy model

Summary A multispecies canopy photosynthesis simulation model was used to examine the importance of canopy structure in influencing light interception and carbon gain in mixed and pure stands of wheat (Triticum aestivum L.) and wild oat (Avena fatua L.), a common weedy competitor of wheat. In the mixtures, the fraction of the simulated canopy photosynthesis contributed by wheat was found to decline during the growing season and this decline was closely related to reductions in the amount of leaf area in upper canopy layers. For both species in mixture and in monoculture, simulated photosynthesis was greatest in the middle or upper-middle canopy layers and sensitivity analyses revealed that canopy photosynthesis was most sensitive to changes in leaf area and leaf inclination in these layers. Changes in LAI and leaf inclination affected canopy carbon gain differently for mixtures and monocultures, but the responses were not the same for the two species. Results from simulations where the structural characteristics of the two species were substituted indicated that species differences in leaf inclination, sheath area and the fraction of leaf area alive were of minor consequence compared with the differences in total leaf area in influencing relative canopy carbon gain in mixtures. Competition for light in these species mixtures appears to be influenced most by differences in the positioning of leaf area in upper canopy layers which determines, to a great extent, the amount of light intercepted.     

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.     

Thermotolerance of Photosystem 2 of Three Alpine Plant Species Under Field Conditions

The species specific response of photosystem 2 (PS2) efficiency and its thermotolerance to diurnal and seasonal alterations in leaf temperature, irradiance, and water relations were investigated under alpine field conditions (1 950 m) and in response to an in situ long-term heat treatment (+3 K). Three plant species were compared using the naturally occurring microstratification of alpine environments, i.e. under contrasting leaf temperatures but under similar macroclimatic conditions. Thermotolerance of PS2 showed a high variability in all three species of up to 9.6 K. Diumal changes (increases or even decreases) in PS2 thermotolerance occurred frequently with a maximum increase of +4.8 K in Loiseleuria procumbens. Increasing leaf temperatures and photosynthetic photon flux density influenced thermotolerance adjustments. Under long-term heating (+3 K) of L. procumbens canopies with infra-red lamps, the maxima of the critical (T_c) and the lethal (T_p) temperature of PS2 increased by at least 1 K. Thermotolerance of the leaf tissue (LT₅₀) increased significantly by +0.6 K. The effects of slight water stress on thermotolerance of PS2 were species specific. High temperature thresholds for photoinhibition were significantly different between species and increased by 9 K from the species in the coldest microhabitat to the species in the warmest. Experimental heating of L. procumbens canopies by +3 K caused a significant (p>0.01) upward shift of the high temperature threshold for photoinhibition by +3 K. Each species appeared to be very well adapted to the thermal conditions of its microhabitat as under the most frequently experienced daytime leaf temperatures no photoinhibition occurred. The observed fine scale thermal adjustment of PS2 in response to increased leaf temperatures shows the potential to optimise photosynthesis under varying environmental conditions as long as the upper thermal limits are not exceeded.     

Temperature dependence of growth,development, and photosynthesis in maize under elevated CO2

Global atmospheric carbon dioxide concentrations (C_a) are rising. As a consequence, recent climate models have projected that global surface air temperature may increase 1.4–5.8 °C with the doubling of C_a by the end of the century. Because, changes in C_a and temperature are likely to occur concomitantly, it is important to evaluate how the temperature dependence of key physiological processes are affected by rising C_a in major crop plants including maize (Zea mays L.), a globally important grain crop with C₄ photosynthetic pathway. We investigated the temperature responses of photosynthesis, growth, and development of maize plants grown at five temperature regimes ranging from 19/13 to 38.5/32.5 °C under current (370 μmol mol⁻¹) and doubled (750 μmol mol⁻¹) C_a throughout the vegetative stages using sunlit controlled environmental chambers in order to test if the temperature dependence of these processes was altered by elevated C_a. Leaf and canopy photosynthetic rates, C₄ enzyme activities, leaf appearance rates, above ground biomass accumulation and leaf area were measured. We then applied temperature response functions (e.g., Arrhenius and Beta distribution models) to fit the measured data in order to provide parameter estimates of the temperature dependence for modeling photosynthesis and development at current and elevated C_a in maize. Biomass, leaf area, leaf appearance rate, and photosynthesis measured at growth C_a was not changed in response to CO₂ enrichment. Carboxylation efficiency and the activities of C₄ enzymes were reduced with CO₂ enrichment indicating possible photosynthetic acclimation of the C₄ cycle. All measured parameters responded to growth temperatures. Leaf appearance rate and leaf photosynthesis showed curvilinear response with optimal temperatures near 32 and 34 °C, respectively. Total above ground biomass and leaf area were negatively correlated with growth temperature. The dependence of leaf appearance rate, biomass, leaf area, leaf and canopy photosynthesis, and C₄ enzyme activities on growth temperatures was comparable between current and elevated C_a. The results of this study suggest that the temperature effects on growth, development, and photosynthesis may remain unchanged in elevated C_a compared with current C_a in maize.     

In situ temperature relationships of biochemical and stomatal controls of photosynthesis in four lowland tropical tree species

Net photosynthetic carbon uptake of Panamanian lowland tropical forest species is typically optimal at 30–32 °C. The processes responsible for the decrease in photosynthesis at higher temperatures are not fully understood for tropical trees. We determined temperature responses of maximum rates of RuBP‐carboxylation (V_CMax) and RuBP‐regeneration (J_Max), stomatal conductance (G_s), and respiration in the light (R_Light) in situ for 4 lowland tropical tree species in Panama. G_s had the lowest temperature optimum (T_Opt), similar to that of net photosynthesis, and photosynthesis became increasingly limited by stomatal conductance as temperature increased. J_Max peaked at 34–37 °C and V_CMax ~2 °C above that, except in the late‐successional species Calophyllum longifolium, in which both peaked at ~33 °C. R_Light significantly increased with increasing temperature, but simulations with a photosynthesis model indicated that this had only a small effect on net photosynthesis. We found no evidence for Rubisco‐activase limitation of photosynthesis. T_Opt of V_CMax and J_Max fell within the observed in situ leaf temperature range, but our study nonetheless suggests that net photosynthesis of tropical trees is more strongly influenced by the indirect effects of high temperature—for example, through elevated vapour pressure deficit and resulting decreases in stomatal conductance—than by direct temperature effects on photosynthetic biochemistry and respiration.     

Carbon balance,productivity, and water use of cold-winter desert shrub communities dominated by C3 and C4 species

Summary Common generalizations concerning the ecologic significance of C₄ photosynthesis were tested in a study of plant gas exchange, productivity, carbon balance, and water use in monospecific communities of C₃ and C₄ salt desert shrubs. Contrary to expectations, few of the hypotheses concerning the performance of C₄ species were supported. Like the C₃ species, Ceratoides lanata, the C₄ shrub, Atriplex confertifolia, initiated growth and photosynthetic activity in the cool spring months and also exhibited maximum photosynthetic rates at this time of year. To compete successfully with C₃ species, Atriplex may have been forced to evolve the capacity for photosynthesis at low temperatures prevalent during the spring when moisture is most abundant. Maximum photosynthetic rates of Atriplex were lower than those of the C₃ species. This was compensated by a prolonged period of low photosynthetic activity in the dry late summer months while Ceratoides became largely inactive. However, the annual photosynthetic carbon fixation per ground area was about the same in these two communities composed of C₃ and C₄ shrubs. The C₄ species did not exhibit greater leaf diffusion resistance than the C₃ species. The photosynthesis/transpiration ratios of the two species were about the same during the period of maximum photosynthetic rates in the spring. During the warm summer months the C₄ species did have superior photosynthesis/transpiration ratios. Yet, since Ceratoides completed a somewhat greater proportion of its annual carbon fixation earlier in the season, the ratio of annual carbon fixation/transpiratory water loss in the two communities was about the same. Atriplex did incorporate a greater percentage of the annual carbon fixation into biomass production than did Ceratoides. However, this is considered to be a reflection of properties apart from the C₄ photosynthetic pathway. Both species displayed a heavy commitment of carbon to the belowground system, and only about half of the annual moisture resource was utilized in both communities.     

Structural and functional variability within the canopyand its relevance for carbon gain and stress avoidance

The functional variability in leaf angle distribution within the canopy was analysed with respect to regulation of light interception and photoprotection. Leaf orientation strongly determined the maximum photochemical efficiency of PSII (F_v/F_m) during summer: horizontal leaves were highly photoinhibited whereas vertical leaf orientation protected the leaves from severe photoinhibition. The importance of leaf orientation within the canopy was analysed in two Mediterranean macchia species with distinct strategies for drought and photoinhibition avoidance during summer. The semi-deciduous species (Cistus monspeliensis) exhibited strong seasonal but minimal spatial variability in leaf orientation. Reversible structural regulation of light interception by vertical leaf orientation during summer protected the leaves from severe photoinhibition. The evergreen sclerophyll (Quercus coccifera) exhibited high spatial variability in leaf angle distribution throughout the year and was less susceptible to photoinhibition. The importance of both strategies for plant primary production was analysed with a three-dimensional canopy photoinhibition model (CANO-PI). Simulations indicated that high variability in leaf angle orientation in Q. coccifera resulted in whole-plant carbon gain during the summer, which was 94 % of the maximum rate achieved by theoretical homogeneous leaf orientations. The high spatial variability in leaf angle orientation may be an effective compromise between efficient light harvesting and avoidance of excessive radiation in evergreen plants and may optimize annual primary production. Whole plant photosynthesis was strongly reduced by water stress and photoinhibition in C. monspeliensis; however, the simulations indicated that growth-related structural regulation of light interception served as an important protection against photoinhibitory reduction in whole-plant carbon gain.     

Effects of elevated temperature on photosynthesis in desert plant <Emphasis Type="Italic">Alhagi sparsifolia</Emphasis> S

Most plants growing in temperate desert zone exhibit brief temperature-induced inhibition of photosynthesis at midday in the summer. Heat stress has been suggested to restrain the photosynthesis of desert plants like Alhagi sparsifolia S. It is therefore possible that high midday temperatures damage photosynthetic tissues, leading to the observed inhibition of photosynthesis. In this study, we investigated the mechanisms underlying heat-induced inhibition of photosynthesis in A. sparsifolia, a dominant species found at the transition zone between oasis and sandy desert on the southern fringe of the Taklamakan desert. The chlorophyll (Chl) a fluorescence induction kinetics and CO₂ response curves were used to analyze the thermodynamic characters of both photosystem II (PSII) and Rubisco after leaves were exposed to heat stress. When the leaves were heated to temperatures below 43°C, the initial fluorescence of the dark-adapted state (F_o), and the maximum photochemical efficiency of PSII (F_v/F_m), the number of active reaction centers per cross section (RCs) and the leaf vitality index (PI) increased or declined moderately. These responses were reversed, however, upon cooling. Moreover, the energy allocation in PSII remained stable. The gradual appearance of a K point in the fluorescence curve at 48°C indicated that higher temperatures strongly impaired PSII and caused irreversible damage. As the leaf temperature increased, the activity of Rubisco first increased to a maximum at 34°C and then decreased as the temperature rose higher. Under high-temperature stress, cell began to accumulate oxidative species, including ammoniacal nitrogen, hydrogen peroxide (H₂O₂), and superoxide (O₂ ^·−), suggesting that disruption of photosynthesis may result from oxidative damage to photosynthetic proteins and thylakoid membranes. Under heat stress, the biosynthesis of nonenzyme radical scavenging carotenoids (Cars) increased. We suggest that although elevated temperature affects the heat-sensitive components comprising of PSII and Rubisco, under moderately high temperature the decrease in photosynthesis is mostly due to inactivation of dark reactions.     

Neglecting acclimation of photosynthesis under drought can cause significant errors in predicting leaf photosynthesis in wheat

Extreme climatic events, such as heat waves, cold snaps and drought spells, related to global climate change, have become more frequent and intense in recent years. Acclimation of plant physiological processes to changes in environmental conditions is a key component of plant adaptation to climate change. We assessed the temperature response of leaf photosynthetic parameters in wheat grown under contrasting water regimes and growth temperatures (T_growth). Two independent experiments were conducted under controlled conditions. In Experiment 1, two wheat genotypes were subjected to well-watered or drought-stressed treatments; in Experiment 2, the two water regimes combined with high, medium and low T_growth were imposed on one genotype. Parameters of a biochemical C₃-photosynthesis model were estimated at six leaf temperatures for each factor combination. Photosynthesis acclimated more to drought than to T_growth. Drought affected photosynthesis by lowering its optimum temperature (T_opt) and the values at T_opt of light-saturated net photosynthesis, stomatal conductance, mesophyll conductance, the maximum rate of electron transport (J_max) and the maximum rate of carboxylation by Rubisco (V_cmax). T_opt for V_cmax was up to 40°C under well-watered conditions but 24–34°C under drought. The decrease in photosynthesis under drought varied among T_growth but was similar between genotypes. The temperature response of photosynthetic quantum yield under drought was partly attributed to photorespiration but more to alternative electron transport. All these changes in biochemical parameters could not be fully explained by the changed leaf nitrogen content. Further model analysis showed that both diffusional and biochemical parameters of photosynthesis and their thermal sensitivity acclimate little to T_growth, but acclimate considerably to drought and the combination of drought and T_growth. The commonly used modelling approaches, which typically consider the response of diffusional parameters, but ignore acclimation responses of biochemical parameters to drought and T_growth, strongly overestimate leaf photosynthesis under variable temperature and drought.     

The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Californian chaparral shrub

The functional roles of the contrasting morphologies of sun and shade shoots of the evergreen shrub Heteromeles arbutifolia were investigated in chaparral and understory habitats by applying a three-dimensional plant architecture simulation model, YPLANT. The simulations were shown to accurately predict the measured frequency distribution of photosynthetic photon flux density (PFD) on both the leaves and a horizontal surface in the open, and gave reasonably good agreement for the more complex light environment in the shade. The sun shoot architecture was orthotropic and characterized by steeply inclined (mean = 71^o) leaves in a spiral phyllotaxy with short internodes. This architecture resulted in relatively low light absorption efficiencies (E _A) for both diffuse and direct PFD, especially during the summer when solar elevation angles were high. Shade shoots were more plagiotropic with longer internodes and a pseudo-distichous phyllotaxis caused by bending of the petioles that positioned the leaves in a nearly horizontal plane (mean = 5^o). This shade-shoot architecture resulted in higher E _A values for both direct and diffuse PFD as compared to those of the sun shoots. Differences in E _A between sun and shade shoots and between summer and winter were related to differences in projection efficiencies as determined by leaf and solar angles, and by differences in self shading resulting from leaf overlap. The leaves exhibited photosynthetic acclimation to the sun and the shade, with the sun leaves having higher photosynthetic capacities per unit area, higher leaf mass per unit area and lower respiration rates per unit area than shade leaves. Despite having 7 times greater available PFD, sun shoots absorbed only 3 times more and had daily carbon gains only double of those of shade shoots. Simulations showed that sun and shade plants performed similarly in the open light environment, but that shade shoots substantially outperformed sun shoots in the shade light environment. The shoot architecture observed in sun plants appears to achieve an efficient compromise between maximizing carbon gain while minimizing the time that the leaf surfaces are exposed to PFDs in excess of those required for light saturation of photosynthesis and therefore potentially photoinhibitory. Received: 8 June 1997 / Accepted: 2 November 1997