植物光合作用限制因素与植被生产力研究进展

随着全球气候变化的加剧,陆地生态系统中植物光合作用限制影响程度的增加已成为降低全球植被净初级生产力的主要因素。系统了解植物光合作用限制因素是科学评估植被生产力的重要前提,也是缓解植物光合作用限制,增加植物光合碳同化能力的先决条件。对植物光合作用限制因素进行了系统解析,分析了光合作用三种限制因素生化限制(Biochemical limitation,l_b)、气孔限制(Stomatal limitation,l_s)、叶肉限制(Mesophyll limitation,l_m)的环境响应,重点讨论了叶肉限制及其影响机理,述评了光合作用限制定量分析方法及改善措施,最后以提高植被生产力为驱使目标,对未来植物光合作用限制因素研究提出以下内容：(1)基因工程技术与系统生物学数据相结合提高植被生产力;(2)气孔响应速度对植物光合作用的影响机制;(3)水通道蛋白(Aquaporin, AQPs)和碳酸酐酶(Carbonic anhydrase, CAs)感知环境信号变化的驱动基因。以期为未来气候变化背景下,深入认识和降低植物光合作用限制,提...     

Down co-regulation of light absorption, photochemistry, and carboxylation in Fe-deficient plants growing in different environments

The regulation of photosynthesis through changes in light absorption, photochemistry, and carboxylation efficiency has been studied in plants grown in different environments. Iron deficiency was induced in sugar beet (Beta vulgaris L.) by growing plants hydroponically in controlled growth chambers in the absence of Fe in the nutrient solution. Pear (Pyrus communis L.) and peach (Prunus persica L. Batsch) trees were grown in field conditions on calcareous soils, in orchards with Fe deficiency-chlorosis. Gas exchange parameters were measured in situ with actual ambient conditions. Iron deficiency decreased photosynthetic and transpiration rates, instantaneous transpiration efficiencies and stomatal conductances, and increased sub-stomatal CO₂ concentrations in the three species investigated. Photosynthesis versus CO₂ sub-stomatal concentration response curves and chlorophyll fluorescence quenching analysis revealed a non-stomatal limitation of photosynthetic rates under Fe deficiency in the three species investigated. Light absorption, photosystem II, and Rubisco carboxylation efficiencies were down-regulated in response to Fe deficiency in a coordinated manner, optimizing the use of the remaining photosynthetic pigments, electron transport carriers, and Rubisco.     

Gas exchange and resource-use efficiency of Leymus cinereus (Poaceae): diurnal and seasonal responses to naturally declining soil moisture

We examined factors that limit diurnal and seasonal photosynthesis in Leymus cinereus, a robust tussock grass from shrub-steppes of western North America. Data from plants in a natural stand and in experimental field plots indicate that this bunchgrass has 1) a high photosynthetic capacity, 2) high leaf nitrogen content and high nitrogen-use efficiency, 3) a steep leaf-to-air diffusion gradient for carbon dioxide, which enhances intrinsic water-use efficiency, and 4) photosynthetic tissues that tolerate severe water stress and recover quickly from moderate water stress. Midday depressions of CO₂ assimilation (A) and stomatal conductance were slight in plants with plentiful water, but marked in plants subject to moderate water stress. Midday stomatal closure in moderately stressed plants reduced intercellular carbon dioxide concentration (c_i) by ≈40 μl liter^-1. The maximum rate of A achieved during the day for severely stressed plants (predawn water potential = -4 MPa) was one-third and daily carbon gain per unit leaf area was about one-fourth that of well-watered plants. For plants in the natural stand, CO₂-saturated photosynthesis declined almost linearly with decreasing soil water availability over the growing season, whereas there was little effect on A at CO₂ ambient levels or on carboxylation efficiency until predawn water potentials reached -1.8 MPa. Nitrogen-use efficiency declined with diminishing soil moisture, but there was no seasonal change in stomatal limitation or instantaneous water-use efficiency as estimated from A vs. c_i curves at optimal leaf temperature and moderate atmospheric evaporative demand. Thus, reduced stomatal conductance in response to increased evaporative demand may increase stomatal limitation diumally, but over the growing season, stomatal limitation estimated from A vs. c_i curves is relatively constant because maximum stomatal conductance is closely tuned to the CO₂ assimilatory capacity of the mesophyll.     

Interactive effects of light,leaf temperature,CO2 and O2 on photosynthesis in soybean

A biochemical model of C ₃photosynthesis has been developed by G.D. Farquhar et al. (1980, Planta 149, 78–90) based on Michaelis-Menten kinetics of ribulose-1,5-bisphosphate (RuBP) carboxylase-oxygenase, with a potential RuBP limitation imposed via the Calvin cycle and rates of electron transport. The model presented here is slightly modified so that parameters may be estimated from whole-leaf gas-exchange measurements. Carbon-dioxide response curves of net photosynthesis obtained using soybean plants (Glycine max (L.) Merr.) at four partial pressures of oxygen and five leaf temperatures are presented, and a method for estimating the kinetic parameters of RuBP carboxylase-oxygenase, as manifested in vivo, is discussed. The kinetic parameters so obtained compare well with kinetic parameters obtained in vitro, and the model fits to the measured data give r ²values ranging from 0.87 to 0.98. In addition, equations developed by J.D. Tenhunen et al. (1976, Oecologia 26, 89–100, 101–109) to describe the light and temperature responses of measured CO₂-saturated photosynthetic rates are applied to data collected on soybean. Combining these equations with those describing the kinetics of RuBP carboxylase-oxygenase allows one to model successfully the interactive effects of incident irradiance, leaf temperature, CO₂ and O₂ on whole-leaf photosynthesis. This analytical model may become a useful tool for plant ecologists interested in comparing photosynthetic responses of different C₃ plants or of a single species grown in contrasting environments.Abbreviations PCO photorespiratory carbon oxidation - PCR photosynthetic carbon reduction - PPFD photosynthetic photon-flux density - RuBP ribulose bisphosphate     

Electron transport is the functional limitation of photosynthesis in field-grown Pima cotton plants at high temperature

Restrictions to photosynthesis can limit plant growth at high temperature in a variety of ways. In addition to increasing photorespiration, moderately high temperatures (35–42 °C) can cause direct injury to the photosynthetic apparatus. Both carbon metabolism and thylakoid reactions have been suggested as the primary site of injury at these temperatures. In the present study this issue was addressed by first characterizing leaf temperature dynamics in Pima cotton (Gossypium barbadense) grown under irrigation in the US desert south‐west. It was found that cotton leaves repeatedly reached temperatures above 40 °C and could fluctuate as much as 8 or 10 °C in a matter of seconds. Laboratory studies revealed a maximum photosynthetic rate at 30–33 °C that declined by 22% at 45 °C. The majority of the inhibition persisted upon return to 30 °C. The mechanism of this limitation was assessed by measuring the response of photosynthesis to CO₂ in the laboratory. The first time a cotton leaf (grown at 30 °C) was exposed to 45 °C, photosynthetic electron transport was stimulated (at high CO₂) because of an increased flux through the photorespiratory pathway. However, upon cooling back to 30 °C, photosynthetic electron transport was inhibited and fell substantially below the level measured before the heat treatment. In the field, the response of assimilation (A) to various internal levels of CO₂ (C_i) revealed that photosynthesis was limited by ribulose‐1,5‐bisphosphate (RuBP) regeneration at normal levels of CO₂ (presumably because of limitations in thylakoid reactions needed to support RuBP regeneration). There was no evidence of a ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) limitation at air levels of CO₂ and at no point on any of 30 A–C_i curves measured on leaves at temperatures from 28 to 39 °C was RuBP regeneration capacity measured to be in substantial excess of the capacity of Rubisco to use RuBP. It is therefore concluded that photosynthesis in field‐grown Pima cotton leaves is functionally limited by photosynthetic electron transport and RuBP regeneration capacity, not Rubisco activity.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Carbon source–sink limitations differ between two species with contrasting growth strategies

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximize crop yield. We investigated how differences in growth rate arise from source–sink limitations, using a model system comparing a fast‐growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow‐growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub‐ambient and elevated CO₂ concentrations ([CO₂]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO₂] indicating that source strength was near maximal at current [CO₂]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO₂] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO₂], and lower non‐structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximizing growth of elite cereals under future elevated [CO₂].     

Does long-term cultivation of saplings under elevated CO2 concentration influence their photosynthetic response to temperature?

Background and Aims Plants growing under elevated atmospheric CO₂ concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO₂ the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase.Methods The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4–5 years to either ambient (AC; 385 µmol mol⁻¹) or elevated (EC; 700 µmol mol⁻¹) CO₂ concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques.Key Results Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO₂ assimilation (A_max) and a decline in photorespiration (R_L) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (T_opt) of A_max, Rubisco carboxylation rates (V_Cmax) and R_L were found for EC saplings compared with AC saplings. However, the shifts in T_opt of A_max were instantaneous, and disappeared when measured at identical CO₂ concentrations. Higher values of T_opt at elevated CO₂ were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry.Conclusions Elevated CO₂ instantaneously increases temperature optima of A_max due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO₂ concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO₂ is unlikely. The hypothesis that long-term cultivation at elevated CO₂ leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary.     

Physiological Impacts of Elevated CO2 Concentration Ranging from Molecular to Whole Plant Responses

The dynamics of the terrestrial ecosystems depend on interactions between a number of biogeochemical cycles (i.e. carbon, nutrient, and hydrological cycles) that may be modified by human actions. Conversely, terrestrial ecosystems are important components of these cycles that create the sources and sinks of important greenhouse gases (e.g. carbon dioxide, methane, nitrous oxide). Especially, carbon is exchanged naturally among these ecosystems and the atmosphere through photosynthesis, respiration, decomposition, and combustion processes. Continuous increase of atmospheric carbon dioxide (CO₂) concentration has led to extensive research over the last two decades, during which more then 1 400 scientific papers describing impacts of elevated [CO₂] (EC) on photosynthesis have been published. However, the degree of response is very variable, depending on species, growing conditions, mineral nutrition, and duration of CO₂ enrichment. In this review, I have summarised the major physiological responses of plants, in particular of trees, to EC including molecular and primary, especially photosynthetic, physiological responses. Likewise, secondary (photosynthate translocation and plant water status) and tertiary whole plant responses including also plant to plant competition are shown.     

Temperature effects on the photosynthetic response of C3 plants to long-term CO2 enrichment

To assess the long-term effect of increased CO₂ and temperature on plants possessing the C₃ photosynthetic pathway, Chenopodium album plants were grown at one of three treatment conditions: (1) 23 °C mean day temperature and a mean ambient partial pressure of CO₂ equal to 350 bar; (2) 34 °C and 350 bar CO₂; and (3) 34 °C and 750 bar CO₂. No effect of the growth treatments was observed on the CO₂ reponse of photosynthesis, the temperature response of photosynthesis, the content of Ribulose-1,5-bisphosphate carboxylase (Rubisco), or the activity of whole chain electron transport when measurements were made under identical conditions. This indicated a lack of photosynthetic acclimation in C. album to the range of temperature and CO₂ used in the growth treatments. Plants from every treatment exhibited similar interactions between temperature and CO₂ on photosynthetic activity. At low CO₂ (< 300 bar), an increase in temperature from 25 to 35 °C was inhibitory for photosynthesis, while at elevated CO₂ (> 400 bar), the same increase in temperature enhanced photosynthesis by up to 40%. In turn, the stimulation of photosynthesis by CO₂ enrichment increased as temperature increased. Rubisco capacity was the primary limitation on photosynthetic activity at low CO₂ (195 bar). As a consequence, the temperature response of A was relatively flat, reflecting a low temperature response of Rubisco at CO₂ levels below its k_m for CO₂. At elevated CO₂ (750 bar), the temperature response of electron transport appeared to control the temperature dependency of photosynthesis above 18 °C. These results indicate that increasing CO₂ and temperature could substantially enhance the carbon gain potential in tropical and subtropical habitats, unless feedbacks at the whole plant or ecosystem level limit the long-term response of photosynthesis to an increase in CO₂ and temperature.Abbreviations A net CO₂ assimilation rate - C _a ambient partial pressure of CO₂ - C _i intercellular partial pressure of CO₂ - Rubisco Ribulose-1,5-bisphosphate carboxylase - VPD vapor pressure difference between leaf and air     

Do open-top chambers overestimate the effects of rising CO2 on plants? An analysis using spring wheat

The microclimate in facilities for studying effects of elevated CO₂ on crops differs from ambient conditions. Open-top chambers (OTCs) increase temperature by 1–3 °C. If temperature and CO₂ interact in their effect on crops, this would limit the value of OTC experiments. Furthermore, interaction of CO₂ and temperature deserves study because increases in atmospheric CO₂ concentration are expected to cause global warming. This paper describes two experiments in which a recently developed cooling system for OTCs was used to analyse the effects of temperature on photosynthesis, growth and yield of spring wheat (Triticum aestivum L., cv. Minaret). Two levels of CO₂ were used (350 and 700 ppm), and two levels of temperature, with cooled OTCs being 1.6–2.4 °C colder than noncooled OTCs. Photosynthetic rates were increased by elevated CO₂, but no effect of temperature was found. Cross-switching CO₂ concentrations as well as determination of A–C_i curves showed that plant photosynthetic capacity after anthesis acclimated to elevated CO₂. The acclimation may be related to the effects of CO₂ on tissue composition: elevated CO₂ decreased leaf nitrogen concentrations and increased sugar content. Calculations of the seasonal mean crop light-use efficiency (LUE) were consistent with the photosynthesis data in that CO₂ increased LUE by 20% on average whereas temperature had no effect. Both elevating CO₂ and cooling increased grain yield, by an average of 11% and 23%, respectively. CO₂ and temperature stimulated yield via different mechanisms: CO₂ increased photosynthetic rate, but decreased crop light interception capacity (LAI), whereas cooling increased grain yield by increasing LAI and extending the growing season with 10 days. The effects of CO₂ and temperature were not additive: the CO₂ effect was about doubled in the noncooled open-top chambers. In most cases, effects on yield were mediated through increased grain density rather than increased individual grain weights. The higher growth response to elevated CO₂ in noncooled vs. cooled OTCs shows that a cooling system may remove a bias towards overestimating crop growth response to CO₂ in open-top chambers.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.     

Improved temperature response functions for models of Rubisco-limited photosynthesis

Predicting the environmental responses of leaf photosynthesis is central to many models of changes in the future global carbon cycle and terrestrial biosphere. The steady-state biochemical model of C₃ photosynthesis of Farquhar et al. (Planta 149, 78–90, 1980) provides a basis for these larger scale predictions; but a weakness in the application of the model as currently parameterized is the inability to accurately predict carbon assimilation at the range of temperatures over which significant photosynthesis occurs in the natural environment. The temperature functions used in this model have been based on in vitro measurements made over a limited temperature range and require several assumptions of in vivo conditions. Since photosynthetic rates are often Rubisco-limited (ribulose, 1-5 bisphosphate carboxylase/oxygenase) under natural steady-state conditions, inaccuracies in the functions predicting Rubisco kinetic properties at different temperatures may cause significant error. In this study, transgenic tobacco containing only 10% normal levels of Rubisco were used to measure Rubisco-limited photosynthesis over a large range of CO₂ concentrations. From the responses of the rate of CO₂ assimilation at a wide range of temperatures, and CO₂ and O₂ concentrations, the temperature functions of Rubisco kinetic properties were estimated in vivo. These differed substantially from previously published functions. These new functions were then used to predict photosynthesis in lemon and found to faithfully mimic the observed pattern of temperature response. There was also a close correspondence with published C₃ photosynthesis temperature responses. The results represent an improved ability to model leaf photosynthesis over a wide range of temperatures (10–40 °C) necessary for predicting carbon uptake by terrestrial C₃ systems.     

Effect of jasmonic acid on the stomatal and nonstomatal limitation of leaf photosynthesis in barley leaves

The effect of long-term (7 days) and shortterm (up to 2 h) treatment of barley plants with jasmonic acid (JA) on the components contributing to stomatal and nonstomatal limitation of photosynthesis was studied. Net CO₂ assimilation rate (A) responses to intercellular CO₂ concentration (C _i ), i.e., A/C _i curves, were used to assess the photosynthetic ability. Long-term treatment of barley plants with JA led to a noticeable decrease in both the initial slope of the A/C _i curves and the maximum A at saturating C _i . The proportion of stomatal and nonstomatal factors in limitation of photosynthesis depended on the applied JA concentration. Short-term treatment with JA affected neither the stomatal conductivity for CO₂ nor the rate of photosynthetic CO₂ assimilation. We suggest that JA may affect photosynthesis indirectly, either as a stress-modulating substance, or through the alterations in gene expression.     

Acclimation of photosynthesis to elevated CO2 and temperature in five British native species of contrasting functional type

Acclimation of photosynthesis to growth at elevated CO₂ concentration varies markedly between species. Species functionally classified as stress-tolerators (S) and ruderals (R), are thought to be incapable, or the least capable, of responding positively in terms of growth to elevated [CO₂]. Is this pattern of response also apparent in leaf photosynthesis of wild S- and R-strategists? Acclimatory loss of a photosynthetic and growth response to elevated [CO₂] is assumed to reflect limitation on capacity to utilize additional photosynthate. The doubling of pre-industrial global [CO₂] is expected to coincide with a 3 °C increase in mean temperature which could stimulate growth; will photosynthetic capacity at elevated [CO₂] be greater when the concurrent temperature increase is simulated? Five species from natural grassland of NW Europe and of contrasting ecological strategy were grown in hemispherical greenhouses, environmentally controlled to track the external microclimate. Within a replicated design, plants were grown at (i) current ambient [CO₂] and temperature, (ii) elevated [CO₂] (ambient + 340 μmol mol^–1) and ambient temperature, (iii) ambient [CO₂] and elevated temperature (ambient + 3 °C), or (iv) elevated [CO₂] and elevated temperature. After 75–104 days, the CO₂ response of light-saturated rates of photosynthesis (A_sat) was analysed in controlled-environment cuvettes in a field laboratory. There was no acclimatory loss of photosynthetic capacity with growth in elevated [CO₂] or elevated temperature over this period in Poa alpina (S), Bellis perennis (R) or Plantago lanceolata (mixed C-S-R strategist), and a significant (P ? ? bl 0.05) increase in capacity in Helianthemum nummularium (S) and Poa annua (R). Photosynthetic rates of leaves grown and measured in elevated [CO₂] were therefore significantly higher than rates for leaves grown and measured in ambient [CO₂], for all species. With the exception of Poa alpina, stomatal conductance and stomatal limitation on A_sat showed no acclimatory response to growth in elevated [CO₂]. Carboxylation efficiency, determined from the initial slope of the response of A_sat to intercellular CO₂ concentration was significantly increased by elevated [CO₂] and elevated temperature in H.nummularium, implying a possible increase in in vivo RubisCO activity. Increased carboxylation efficiency of this species was also reflected by an increase in the CO₂- and light-saturated rates of photosynthesis, indicating an increased capacity for regeneration of the primary CO₂ acceptor in photosynthesis. The results show that R-strategists and slow-growing S-strategists, are inherently capable of large increases in leaf photosynthetic capacity with growth in elevated [CO₂] in contrast to expectations from growth studies. With the exception of P.annua, where there was a significant negative interaction between CO₂ and temperature, concurrent increase in growth temperature had little effect on this pattern of response.     

Effects of low and elevated CO2 on C3 and C4 annuals

Abutilon theophrasti (C₃) and Amaranthus retroflexus (C₄), were grown from seed at four partial pressures of CO₂: 15 Pa (below Pleistocene minimum), 27 Pa (pre-industrial), 35 Pa (current), and 70 Pa (future) in the Duke Phytotron under high light, high nutrient, and wellwatered conditions to evaluate their photosynthetic response to historic and future levels of CO₂. Net photosynthesis at growth CO₂ partial pressures increased with increasing CO₂ for C₃ plants, but not C₄ plants. Net photosynthesis of Abutilon at 15 Pa CO₂ was 70% less than that of plants grown at 35 Pa CO₂, due to greater stomatal and biochemical limitations at 15 Pa CO₂. Relative stomatal limitation (RSL) of Abutilon at 15 Pa CO₂ was nearly 3 times greater than at 35 Pa CO₂. A photosynthesis model was used to estimate ribulose-1,5-bisphosphate carboxylase (rubisco) activity (Vc_max), electron transport mediated RuBP regeneration capacity (J _max), and phosphate regeneration capacity (PiRC) in Abutilon from net photosynthesis versus intercellular CO₂ (A–C _i) curves. All three component processes decreased by approximately 25% in Abutilon grown at 15 Pa compared with 35 Pa CO₂. Abutilon grown at 15 Pa CO₂ had significant reductions in total rubisco activity (25%), rubisco content (30%), activation state (29%), chlorophyll content (39%), N content (32%), and starch content (68%) compared with plants grown at 35 Pa CO₂. Greater allocation to rubisco relative to light reaction components and concomitant decreases in J _max and PiRC suggest co-regulation of biochemical processes occurred in Abutilon grown at 15 Pa CO₂. There were no significant differences in photosynthesis or leaf properties in Abutilon grown at 27 Pa CO₂ compared with 35 Pa CO₂, suggesting that the rise in CO₂ since the beginning of the industrial age has had little effect on the photosynthetic performance of Abutilon. For Amaranthus, limitations of photosynthesis were balanced between stomatal and biochemical factors such that net photosynthesis was similar in all CO₂ treatments. Differences in photosynthetic response to growth over a wide range of CO₂ partial pressures suggest changes in the relative performance of C₃ and C₄ annuals as atmospheric CO₂ has fluctuated over geologic time.     

贵州喀斯特森林三种植物对不同坡位环境的光合生理响应

该研究以贵州普定喀斯特森林中、下坡位生长的构树( Broussonetia papyrifera)、朴树( Celtis sinensis)和光滑悬钩子( Rubus tsangii)为材料,通过对碳酸酐酶( CA)活性、光合作用日变化、净光合速率对CO2与光的响应曲线、叶绿素荧光特性以及稳定碳同位素组成等指标的测定,进而对比分析三种植物不同的光合生理响应特性。结果表明：构树光合作用过程的无机碳源既可来自大气中的CO2,也可以在气孔部分闭合的情况下利用细胞内的HCO3－,下坡位的构树较高的CA活性使其利用HCO3－的效率会更高,并能在较低光强下具有较高的光能利用效率。这可能与下坡位的构树具有较高的CA活性有关,对下坡位具有更好的适应性。朴树光合无机碳的同化能力最低,且光合无机碳源较单一,主要利用大气CO2,其较慢的生长速率使其对无机碳的需求最低,且能保持较稳定的无机碳同化速率。相对来说,中坡位的朴树具有相对较高的净光合速率和光能利用效率,对中坡位表现出较好的适应性。光滑悬钩子主要利用大气中的CO2进行光合作用。中坡位的光滑悬钩子具有较强的光能利用效率,并表现出较高的净光合速率,光滑悬钩子对中坡位同样表现出较好的适应性。该研究结果为喀斯特生态脆弱区植被重建过程中树种的选择及合理配置提供了科学依据。     

CO2 alters water use,carbon gain,and yield for the dominant species in a natural grassland

Global atmospheric CO₂ is increasing at a rate of 1.5–2 ppm per year and is predicted to double by the end of the next century. Understanding how terrestrial ecosystems will respond in this changing environment is an important goal of current research. Here we present results from a field study of elevated CO₂ in a California annual grassland. Elevated CO₂ led to lower leaf-level stomatal conductance and transpiration (approximately 50%) and higher mid-day leaf water potentials (30–35%) in the most abundant species of the grassland, Avena barbata Brot. Higher CO₂ concentrations also resulted in greater midday photosynthetic rates (70% on average). The effects of CO₂ on stomatal conductance and leaf water potential decreased towards the end of the growing season, when Avena began to show signs of senescence. Water-use efficiency was approximately doubled in elevated CO₂, as estimated by instantaneous gas-exchange measurements and seasonal carbon isotope discrimination. Increases in CO₂ and photosynthesis resulted in more seeds per plant (30%) and taller and heavier plants (27% and 41%, respectively). Elevated CO₂ also reduced seed N concentrations (9%).     

Photosynthetic capacity and temperature responses of photosynthesis of rubber trees (Hevea brasiliensis Müll. Arg.) acclimate to changes in ambient temperatures

The aim of this study was to assess the temperature response of photosynthesis in rubber trees (Hevea brasiliensis Müll. Arg.) to provide data for process-based growth modeling, and to test whether photosynthetic capacity and temperature response of photosynthesis acclimates to changes in ambient temperature. Net CO₂ assimilation rate (A) was measured in rubber saplings grown in a nursery or in growth chambers at 18 and 28°C. The temperature response of A was measured from 9 to 45°C and the data were fitted to an empirical model. Photosynthetic capacity (maximal carboxylation rate, V _cmax, and maximal light driven electron flux, J _max) of plants acclimated to 18 and 28°C were estimated by fitting a biochemical photosynthesis model to the CO₂ response curves (A–C _i curves) at six temperatures: 15, 22, 28, 32, 36 and 40°C. The optimal temperature for A (T _opt) was much lower in plants grown at 18°C compared to 28°C and nursery. Net CO₂ assimilation rate at optimal temperature (A _opt), V _cmax and J _max at a reference temperature of 25°C (V _cmax25 and J _max25) as well as activation energy of V _cmax and J _max (E _aV and E _aJ) decreased in individuals acclimated to 18°C. The optimal temperature for V _cmax and J _max could not be clearly defined from our response curves, as they always were above 36°C and not far from 40°C. The ratio J _max25/V _cmax25 was larger in plants acclimated to 18°C. Less nitrogen was present and photosynthetic nitrogen use efficiency (V _cmax25/N _a) was smaller in leaves acclimated to 18°C. These results indicate that rubber saplings acclimated their photosynthetic characteristics in response to growth temperature, and that higher temperatures resulted in an enhanced photosynthetic capacity in the leaves, as well as larger activation energy for photosynthesis.     

Canopy warming caused photosynthetic acclimation and reduced seed yield in maize grown at ambient and elevated [CO2]

Rising atmospheric CO₂ concentration ([CO₂]) and attendant increases in growing season temperature are expected to be the most important global change factors impacting production agriculture. Although maize is the most highly produced crop worldwide, few studies have evaluated the interactive effects of elevated [CO₂] and temperature on its photosynthetic physiology, agronomic traits or biomass, and seed yield under open field conditions. This study investigates the effects of rising [CO₂] and warmer temperature, independently and in combination, on maize grown in the field throughout a full growing season. Free‐air CO₂ enrichment (FACE) technology was used to target atmospheric [CO₂] to 200 μmol mol^?1 above ambient [CO₂] and infrared heaters to target a plant canopy increase of 3.5 °C, with actual season mean heating of ~2.7 °C, mimicking conditions predicted by the second half of this century. Photosynthetic gas‐exchange parameters, leaf nitrogen and carbon content, leaf water potential components, and developmental measurements were collected throughout the season, and biomass and yield were measured at the end of the growing season. As predicted for a C₄ plant, elevated [CO₂] did not stimulate photosynthesis, biomass, or yield. Canopy warming caused a large shift in aboveground allocation by stimulating season‐long vegetative biomass and decreasing reproductive biomass accumulation at both CO₂ concentrations, resulting in decreased harvest index. Warming caused a reduction in photosynthesis due to down‐regulation of photosynthetic biochemical parameters and the decrease in the electron transport rate. The reduction in seed yield with warming was driven by reduced photosynthetic capacity and by a shift in aboveground carbon allocation away from reproduction. This field study portends that future warming will reduce yield in maize, and this will not be mitigated by higher atmospheric [CO₂] unless appropriate adaptation traits can be introduced into future cultivars.