Stomatal patchiness and task-performing networks

BACKGROUND: Patchy stomatal conductance is a poorly understood and little-studied phenomenon. It is relatively common, yet it appears to be detrimental to water-use efficiency under some conditions and has no immediately obvious physiological function of any kind. Much of the difficulty in studying patchy stomatal conductance is tied to its unpredictability, both in occurrence and in characteristics. SCOPE AND CONCLUSIONS: Statistical analyses of the variability of stomatal patchiness reveal remarkable similarities to structures and behaviours found in locally connected networks of dynamic units that perform tasks. Such systems solve problems that reside at the level of the entire network despite the absence of a central processor or a mechanism for directly sharing information over the entire system. Frequently, task performance is emergent, in the sense that no unit independently performs the task. Because each unit in the network can communicate with only its immediate neighbours, problem solving is accomplished by the states of the individual units self-organizing into synchronized, collective patterns. In some cases, patches of states form and move coherently over the network, thus providing a means for distantly separated parts of the network to communicate. Often, exactly what form these patches take and how they move as the units synchronize is highly unpredictable. In analogy with such networks, it is suggested that stomatal patchiness may be a signature that plants optimize gas exchange in a more sophisticated and adaptive manner than if performed by their individual stomata independently.     

Coordination of leaf and stem water transport properties in tropical forest trees

Stomatal regulation of transpiration constrains leaf water potential (Ψ_L) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the stems to which they are attached. We evaluated a suite of leaf and stem functional traits governing water relations in individuals of 11 lowland tropical forest tree species to determine the manner in which the traits were coordinated with stem xylem vulnerability to embolism. Stomatal regulation of Ψ_L was associated with minimum values of water potential in branches (Ψ_br) whose functional significance was similar across species. Minimum values of Ψ_br coincided with the bulk sapwood tissue osmotic potential at zero turgor derived from pressure–volume curves and with the transition from a linear to exponential increase in xylem embolism with increasing sapwood water deficits. Branch xylem pressure corresponding to 50% loss of hydraulic conductivity (P ₅₀) declined linearly with daily minimum Ψ_br in a manner that caused the difference between Ψ_br and P ₅₀ to increase from 0.4 MPa in the species with the least negative Ψ_br to 1.2 MPa in the species with the most negative Ψ_br. Both branch P ₅₀ and minimum Ψ_br increased linearly with sapwood capacitance (C) such that the difference between Ψ_br and P ₅₀, an estimate of the safety margin for avoiding runaway embolism, decreased with increasing sapwood C. The results implied a trade-off between maximizing water transport and minimizing the risk of xylem embolism, suggesting a prominent role for the buffering effect of C in preserving the integrity of xylem water transport. At the whole-tree level, discharge and recharge of internal C appeared to generate variations in apparent leaf-specific conductance to which stomata respond dynamically.     

Disruption of a gene encoding C4-dicarboxylate transporter-like protein increases ozone sensitivity through deregulation of the stomatal response in Arabidopsis thaliana

To understand better the plant response to ozone, we isolated and characterized an ozone-sensitive (ozs1) mutant strain from a set of T-DNA-tagged Arabidopsis thaliana ecotype Columbia. The mutant plants show enhanced sensitivity to ozone, desiccation and sulfur dioxide, but have normal sensitivity to hydrogen peroxide, low temperature and high light levels. The T-DNA was inserted at a single locus which is linked to ozone sensitivity. Identification of the genomic sequences flanking the T-DNA insertion revealed disruption of a gene encoding a transporter-like protein of the tellurite resistance/C(4)-dicarboxylate transporter family. Plants with either of two different T-DNA insertions in this gene were also sensitive to ozone, and these plants failed to complement ozs1. Transpiration levels, stomatal conductance levels and the size of stomatal apertures were greater in ozs1 mutant plants than in the wild type. The stomatal apertures of ozs1 mutant plants responded to light fluctuations but were always larger than those of the wild-type plants under the same conditions. The stomata of the mutant and wild-type plants responded similarly to stimuli such as light, abscisic acid, high concentrations of carbon dioxide and ozone. These results suggest that OZS1 helps to close stomata, being not involved in the responses to these signals.     

Reconstructing atmospheric carbon dioxide with stomata: possibilities and limitations of a botanical pCO2-sensor

Stomatal frequency is often observed to vary inversely with atmospheric CO₂ concentration (pCO₂). The response is due to (1) individual phenotypic plasticity and (2) evolutionary change, depending on the time scale. Evolutionary responses occur more frequently than individual responses and individual responses are more pronounced under subambient pCO₂ levels than under elevated pCO₂ (CO₂ ceiling). The evolutionary response appears therefore to be a valuable device for determining past pCO₂. Since tree leaves often represent a conspicuous and rich resource of fossil material, they are increasingly important in this respect. Additionally, certain tree species are considered to represent living fossils and therefore valuable sources of ancient stomatal data. There are, however, numerous difficulties which have to be considered such as: (1) high variance of the data, especially for fossil material, (2) interspecific differences of the response, (3) the CO₂ ceiling and (4) differences between short-term and long-term responses. Whereas the qualitative pCO₂ signal of stomatal frequency appears to be reliable, quantitative pCO₂ reconstruction has to be performed with caution. The results of a number of studies which used stomatal frequency as a pCO₂ sensor demonstrate good agreement with the results obtained with other proxy data. Current techniques are based on transfer functions which calibrate the fossil data with extant material. It is suggested that a mechanistic approach including physical as well as physiological processes could improve pCO₂ reconstruction. Furthermore, the topic of the influence of pCO₂ on stomatal frequency is significant not only for reconstructing past pCO₂ but also with respect to the climate-biosphere interrelationship.     

弱光下生长的葡萄叶片蒸腾速率和气孔结构的变化

 植物能够对生长环境产生生态适应性,这种适应性可从气孔导度、光合速率、水分利用效率等生态指标上反映出来。为了研究葡萄蒸腾特性对弱光环境的适应性变化,本试验以‘京玉’葡萄幼苗（Vitis vinefera cv. Jingyu）为试验材料,通过遮光处理（2个处理,分别遮光65%和85%）营造弱光环境,测定了在弱光环境下生长的葡萄叶片蒸腾速率、气孔导度、水分利用效率对光照强度的响应,同时用扫描电镜技术观察了气孔的发育。结果表明,弱光环境下生长的葡萄幼苗,叶片的水势较高,但水分利用效率较低,叶片蒸腾速率和气孔导度变化对光照强度的响应缓慢,而自然光下生长的葡萄叶片则反应较迅速。通过对气孔结构的研究发现,与自然光照环境下生长的植株相比,在弱光环境下生长的葡萄幼苗,叶片下表皮的气孔横轴变宽,大小气孔之间差异减少,气孔外突,表皮细胞变大甚至扭曲,角质层变薄。说明葡萄幼苗能够对弱光环境产生适应性变化,其蒸腾特性的变化与其气孔结构的变化相关,具有一致性。     

13C discrimination during CO2 assimilation by the terrestrial biosphere

Estimates of the extent of the discrimination against¹³CO₂ during photosynthesis (_A) on a global basis were made using gridded data sets of temperature, precipitation, elevation, humidity and vegetation type. Stomatal responses to leaf-to-air vapour mole fraction difference (D, leaf-to-air vapour pressure difference divided by atmospheric pressure) were first determined by a literature review and by assuming that stomatal behaviour results in the optimisation of plant water use in relation to carbon gain. Using monthly time steps, modelled stomatal responses toD were used to calculate the ratio of stomatal cavity to ambient CO₂ mole fractions and then, in association with leaf internal conductances, to calculate _A. Weighted according to gross primary productivity (GPP, annual net CO₂ asimilation per unit ground area), estimated _A for C₃ biomes ranged from 12.9 for xerophytic woods and shrub to 19.6 for cool/cold deciduous forest, with an average value from C₃ plants of 17.8. This is slightly less than the commonly used values of 18–20. For C₄ plants the average modelled discrimination was 3.6, again slightly less than would be calculated from C₄ plant dry matter carbon isotopic composition (yielding around 5). From our model we estimate that, on a global basis, 21% of GPP is by C₄ plants and for the terrestrial biosphere as a whole we calculate an average isotope discrimination during photosynthesis of 14.8. There are large variations in _A across the globe, the largest of which are associated with the precence or absence of C₄ plants. Due to longitudinal variations in _A, there are problems in using latitudinally averaged terrestrial carbon isotope discriminations to calculate the ratio of net oceanic to net terrestrial carbon fluxes.     

The role of the epidermis in the generation of the circadian rhythm of carbon dioxide fixation in leaves of Bryophyllum fedtschenkoi

The role of the epidermis in the generation of the endogenous circadian rhythm of CO₂ exchange in leaves of Bryophyllum fedtschenkoi has been examined. At 25° C the rhythm of CO₂ output exhibited by whole leaves kept in continuous darkness and an initially CO₂-free air stream also occurs in isolated pieces of mesophyll. The sensitivity to light of the rhythms in whole leaves and in isolated mesophyll appears to be identical. At 15° C, however, no rhythm is observed in isolated mesophyll tissue, despite there being a conspicuous rhythm in intact leaves. The rhythm of net CO₂ assimilation in whole leaves kept in continuous light and a stream of normal air at either 25° C or at 15° C is abolished by removal of the epidermis, although at 15° C and under the higher of the two light levels used, there is an indication that rhythmicity may begin to reappear after the third day of the experiment. Thus, only under certain environmental conditions is the rhythm of CO₂ exchange in Bryophyllum leaves independent of the epidermis. The results indicate that the rhythm of carbon dioxide fixation in continuous darkness and CO₂-free air is generated primarily in the mesophyll cells, whereas the rhythm in continuous light and normal air is generated in the stomatal guard cells or in an interaction of these cells with the mesophyll cells.Abbreviation PEPCase phosphoenolpyruvate carboxylase     

Carbon-water balance and patchy stomatal conductance

Stomata govern carbon-water balance by simultaneously controlling photosynthesis (A) and transpiration (E). It is unclear how patchy stomatal conductance influences this control. Cowan and Farquhar showed that for a given water supply available during a fixed time interval, carbon gain is maximized by a pattern of stomatal behavior that keeps the partial derivative of A with respect to E constant. This result implies that spatially uniform stomatal conductance is optimal (provided photosynthetic performance and environmental conditions are spatially uniform), so patchy stomatal conductance should be detrimental to carbon-water balance. However, these results required that the curvature of A versus E be uniformly negative. Using mathematical arguments and computer modeling, we show that (1) this caveat is violated under some environmental conditions, (2) water-use efficiency (A/E) is nearly unaffected, and can actually be improved, by patchiness under these conditions, and (3) patchiness has most often been observed under conditions similar to these. These results imply that under many conditions, patchiness may not significantly influence carbon-water balance, consistent with recent work suggesting patchiness may be common but unobserved. Additionally, we discuss implications of these results that muddle the definition of `optimal' in the context of plant gas exchange in some situations, and extend the work of Cowan and Farquhar under conditions causing positive curvature in A versus E. Received: 15 May 1998 / Accepted: 14 October 1998     

湿度对葡萄,樱桃离体繁殖植株气孔行为和移栽成活率的影响

本文通过葡萄和樱桃不同品种离体植株气孔在繁殖不同时期对多种关闭诱导因素的反应,研究了气孔不能关闭的原因和提高植株移栽成活率的影响因素。材料和方法选用二倍     

Analysing the responses of photosynthetic CO2 assimilation to long-term elevation of atmospheric CO2 concentration

Understanding how photosynthetic capacity acclimatises when plants are grown in an atmosphere of rising CO₂ concentrations will be vital to the development of mechanistic models of the response of plant productivity to global environmental change. A limitation to the study of acclimatisation is the small amount of material that may be destructively harvested from long-term studies of the effects of elevation of CO₂ concentration. Technological developments in the measurement of gas exchange, fluorescence and absorption spectroscopy, coupled with theoretical developments in the interpretation of measured values now allow detailed analyses of limitations to photosynthesisin vivo. The use of leaf chambers with Ulbricht integrating spheres allows separation of change in the maximum efficiency of energy transduction in the assimilation of CO₂ from changes in tissue absorptance. Analysis of the response of CO₂ assimilation to intercellular CO₂ concentration allows quantitative determination of the limitation imposed by stomata, carboxylation efficiency, and the rate of regeneration of ribulose 1:5 bisphosphate. Chlorophyll fluorescence provides a rapid method for detecting photoinhibition in heterogeneously illuminated leaves within canopies in the field. Modulated fluorescence and absorption spectroscopy allow parallel measurements of the efficiency of light utilisation in electron transport through photosystems I and IIin situ.Abbreviations A net rate of CO₂ uptke per unit leaf area (µmol m^–2 s^–1) - A_sat light-saturated A - A₈₂₀ change in absorptance of PSI on removal of illumination (OD) - c CO₂ concentration in air (µmol mol^–1) - c_a c in the bulk air; c_i, c in the intercellular spaces - ce carboxylation efficiency (mol m^–2 s^–1) - E transpiration per unit leaf area (mol m^–2 s^–1) - F fluorescence emission of PSII (relative units) - F_m maximal level of F - F_o minimal level of F upon illumination when PSII is maximally oxidised - F_s the steady-state F following the m peak - F_v the difference between F_m and F_o - F'_m maximal F' generated after the m peak by addition of a saturating light pulse - F'_o the minimal level of F' after the m peak determined by re-oxidising PSII by far-red light - g₁ leaf conductance to CO₂ diffusion in the gas phase (mol m^–2 s^–1) - g'₁ leaf conductance to water vapour diffusion in the gas phase (mol m^–2 s^–1) - k_c and k_o the Michaelis constants for CO₂ and O₂, respectively, (µmol mol^–1); - J_max the maximum rate of regeneration of rubP (µmol m^–2 s^–1) - l stomatal limitation to CO₂ uptake (dimensionless, 0–1) - LCP light compensation point of photosynthesis (µmol m^–2 s^–1) - o_i the intercellular O₂ concentration (mmol mol^–1) - Pi cytosol inorganic phosphate concentration - PSI photosystem I - PSII photosystem II - Q photon flux (µmol m^–2 s^–1) - Q_abs Q absorbed by the leaf - rubisCO ribulose 1:5 bisphosphate carboxylase/oxygenase; rubP, ribulose 1:5 bisphosphate; s, projected surface area of a leaf (m²) - V_c,max is the maximum rate of carboxylation (µmol m^–2 s^–1) - W_c the rubisCO limited rate of carboxylation (µmol m^–2 s¹) - W_j the electron transport limited rate of regeneration of rubP (µmol m^–2 s^–1) - W_p the inorganic phosphate limited rate of regeneration of rubP (µmol m^–2 s^–1) - absorptance of light (dimensionless, 0–1) - _a of standard black absorber ₁, of leaf - _s of integrating sphere walls - , CO₂ compensation point of photosynthesis (µmol mol^–1) - the specificity factor for rubisCO carboxylation (dimensionless) - , convexity of the response of A to Q (dimensionless 0–1) - the quantum yield of photosynthesis on an absorbed light basis (A/Q_abs; dimensionless) - the quantum yield of photosynthesis on an incident light basis (A/Q; dimensionless) - _app the maximum - _m the maximum - _m,app the photochemical efficiency of PSII (dimensionless, 0–1) - _PSII,m the maximum