Tansley Review No. 59 Leaf boundary layers

Studies of heat and mass exchange between leaves and their local environment are central to our understanding of plant-atmosphere interactions. The transfer across aerodynamic leaf boundary layers is generally described by non-dimensional expressions which reflect largely empirical adaptations of engineering models derived for flat plates. This paper reviews studies on leaves, and leaf models with varying degrees of abstraction, in free and forced convection. It discusses implecations of finding for leaf morphology as it affects – and is affected by – the local microclimate. Predictions of transfer from many leaves in plant communities are complicated by physical and physiological feedback mechanisms between leaves and their environment. Some common approaches, and the current challenge of integrating leaf-atmosphere interactions into models of global relevance, are also briefly addressed.     

Effects of short-term salinity on leaf gas exchange of the fig (Ficus carica L.)

The influence of short-term salinity (day 1–day 2: 50 mol m^–3 NaCl, day 3–day 7: 100 mol m^–3 NaCl in the nutrient solution) on leaf gas exchange characteristics were studied in two fig clones (Ficus carica L.), whose root mass had been varied in relation to the leaf area. The stomatal conductance was diminished by NaCl in the first week of treatment. NaCl slightly reduced the calculated intercellular partial pressure of CO₂. The net photosynthetic rate of plants with many roots was stimulated by NaCl on some days of the first week of treatment, whereas the net assimilation rate of the plants with few roots remained unaltered or decreased by NaCl. Only the assimilation of the salt-treated plants of one clone for some days during the first week of treatment seemed to be influenced by stomatal conductance. Nonstomatal factors were primarily responsible for the changes in CO₂ uptake in response to salt and/or root treatment. The water use efficiency increased during several days of the first week of NaCl treatment. Decreased stomatal conductance, increased water use efficiency and stimualtion of the net CO₂ assimilation rate appear to enhance salt tolerance during the first few days of salinity. ei]H Lambers     

Stomatal numbers of soybean and response to water stress

The relationship among stomatal density, photosynthetic rate, leaf conductance, plant growth, bean yield and kaempferol triglucoside (K9) in the leaves of soybean (Glycine max (L.) Merr.) was examined in two field tests. K9 in the leaves was associated with reduced stomatal density, reduced photosynthetic rate, reduced stomatal conductance, reduced plant weight and lower bean yield. Plants with high stomatal frequency (lacking K9) were better able to take advantage of increased water supply by increasing stomatal conductance (upper surface), transpiration and bean yield. Plants with low stomatal frequency (with K9) were unresponsive to irrigation and in this sense were more tolerant of water stress, but their overall yield was low.     

Heterogeneity of soil and soil solution chemistry under Norway spruce (Picea abies Karst.) and European beech (Fagus silvatica L.) as influenced by distance from the stem basis

The present study aims at characterizing plant water status under field conditions on a daily basis, in order to improve operational predictions of plant water stress. Ohm's law analog serves as a basis for establishing daily soil-plant relationships, using experimental data from a water-limited soybean crop: 227-1. The daily transpiration flux, T, is estimated from experimental evapotranspiration data and simulated soil evaporation values. The difference, 227-2, named the effective potential gradient, is derived from i) the midday leaf potential of the uppermost expanded leaves and ii) an effective soil potential accounting for soil potential profile and an effectiveness factor of roots competing for water uptake. This factor is experimentally estimated from field observation of roots. G is an apparent hydraulic conductance of water flow from the soil to the leaves. The value of the lower potential limit for water extraction, required to assess the effective soil potential, is calculated with respect to the plant using the predawn leaf potential. It is found to be equal to –1.2 MPa. It appears that over the range of soil and climatic conditions experienced, the daily effective potential gradient remains constant (1.2 MPa), implying that, on a daily basis, transpiration only depends on the hydraulic conductance. The authors explain this behaviour by diurnal variation of osmotic potential, relying on Morgan's theory (1984). Possible generalization of the results to other crop species is suggested, providing a framework for reasoning plant water behaviour at a daily time step.     

Leaf responses to soil water deficits: Comparative sensitivity of leaf expansion rate and leaf conductance in field-grown sunflower (Helianthus annuus L.)

The relative importance of changes in leaf expansion rate (LER) and leaf conductance (g₁) in the control of crop transpiration depends primarily on their sensitivity to soil water deficits. The aim of this paper was to quantify the responses of LER and g₁ to soil water deficits in sunflower (Helianthus annuus L.) under conditions of moderate (spring) and high (summer) evaporative demand. Soil water content, g₁, and LER were measured in dryland (DRY) and daily-irrigated (WET) crops established on a deep sandy-loam (Typic Xerofluvent) in a Mediterranean environment. There was no difference between g₁ of DRY and WET plants (p>0.20) in contrast with a highly significant difference in LER (p<0.001). Even under the harsh conditions of the summer experiment, g₁ did not respond to water deficit in a ten-day period in which LER of DRY plants was reduced to approx. 30% of that measured in WET controls. This field study indicates that g₁ plays at most a minor role in the control of sunflower transpiration in the pre-anthesis period and confirms the importance of leaf expansion in the regulation of gas exchange of expanding canopies subjected to soil water deficits.     

Estimating transpiration from 6-year-old Eucalyptus grandis trees: development of a canopy conductance model and comparison with independent sap flux measurements

Daily patterns of stomatal conductance (g_s), xylem pressure potential (P) and canopy microclimatic variables were recorded on 11 sample days as part of a one-year study of the water use of Eucalyptus grandis Hill ex Maiden in the eastern Transvaal, South Africa. Measured g_s was found to be largely controlled by quantum flux density (Q) and ambient vapour pressure deficit (D). Canopy conductance (g_c) was determined for hourly intervals using g_s measurements and leaf areas in four different canopy levels. A simple model was constructed to allow the prediction of g_c and transpiration from Q, D and season of year. The model was used to estimate transpiration rates from 10 trees in a later study of similarly-aged E. grandis trees, in which sap flow in each tree was measured using the heat pulse velocity (HPV) technique. Five of the trees were monitored on a summer day and five on a winter day. Correspondence between HPV sap flow and modelled transpiration was good for the summertime comparisons, but measured winter-time sap flow rates were underestimated by the model, especially under conditions of high sap flow. The discrepancy is believed to result from having insufficient data from the conductance study to describe the response of g_s to relatively high D in winter. Marked variation in transpiration per unit leaf area indicates that a relatively large number of trees must be sampled for the HPV technique to be used to obtain a mean rate for an entire stand in winter.     

Simulation of the light-induced oscillations of the membrane potential in Potamogeton leaf cells

Summary An attempt has been made to simulate the light-induced oscillations of the membrane potential of Potamogeton lucens leaf cells in relation to the apoplastic pH changes. Previously it was demonstrated that the membrane potential of these cells can be described in terms of proton movements only. It is hypothesized that the membrane potential is determined by an electrogenic H⁺-ATPase with a variable H⁺/ATP stoichiometry. The stoichiometry shifts from a value of two in the dark to a value of one in the light. Moreover, this H⁺ pump shows the characteristics of either a pump or a passive H⁺ conductance: the mode of operation of the H⁺ translocator is considered to be regulated by the external pH. The pump conductance is assumed to be dominant at low or neutral pH, while the passive H⁺ conductance becomes more significant at alkaline pH. The pH dependence of the transport characteristic is expressed by protonation reactions in the plasma membrane. The proposed model can account for most features of the light-induced oscillations but not for the absolute level of the membrane potential.This research was supported by the Foundation of Biophysics, part of the Dutch Organization for Scientific Research (NWO) ECOTRANS publication No. 34.     

Hyposensitive canopy conductance renders ecosystems vulnerable to meteorological droughts

Increased meteorological drought intensity with rising atmospheric demand for water (hereafter vapor pressure deficit [VPD]) increases the risk of tree mortality and ecosystem dysfunction worldwide. Ecosystem-scale water-use strategy is increasingly recognized as a key factor in regulating drought-related ecosystem responses. However, the link between water-use strategy and ecosystem vulnerability to meteorological droughts is poorly established. Using the global flux observations, historic hydroclimatic data, remote-sensing products, and plant functional-trait archive, we identified potentially vulnerable ecosystems, examining how ecosystem water-use strategy, quantified by the percentage bias (δ) of the empirical canopy conductance sensitivity to VPD relative to the theoretical value, mediated ecosystem responses to droughts. We found that prevailing soil water availability substantially impacted δ in dryland regions where ecosystems with insufficient soil moisture usually showed conservative water-use strategy, while ecosystems in humid regions exhibited more pronounced climatic adaptability. Hyposensitive and hypersensitive ecosystems, classified based on δ falling below or above the theoretical sensitivity, respectively, achieved similar net ecosystem productivity during droughts, employing different structural and functional strategies. However, hyposensitive ecosystems, risking their hydraulic system with a permissive water-use strategy, were unable to recover from droughts as quickly as hypersensitive ones. Our findings highlight that processed-based models predicting current functions and future performance of vegetation should account for the greater vulnerability of hyposensitive ecosystems to intensifying atmospheric and soil droughts.     

Different ionic conductances are modulated during the late receptor potential and the prolonged depolarizing afterpotential in Hermissenda type A photoreceptors

Wavelength-dependent, bistable phenomena were found in the receptor potential of Hermissenda crassicornis type A photoreceptors. Short exposure to blue light induced a prolonged depolarizing afterpotential (PDA) following the cessation of the light stimulus. Stronger adaptation to blue light, as caused by prolonged exposure and/or high intensity stimulation, effected a reduction in the early depolarizing transient of the late receptor potential (LRP) as elicited by subsequent stimuli. Vast separation of LRP emergence and PDA emergence could be obtained in photoreceptors in which a strong cancellation of the LRP was accomplished but a PDA still emerged after cessation of the light stimulus. Short exposure to yellow light cancelled the PDA, and stronger adaptation restored the LRP (opposite effect to blue light). The initial depolarizing part of the LRP had earlier been demonstrated to be mediated by the lightdependent increase of an inward conductance. In contrast, in this study the PDA was found to be accompanied by the reduction of an outward conductance, most likely a K⁺ conductance. A bistable photopigment system is thought to control the bistable receptor potential phenomenology by regulating the different membrane conductances during the LRP and the PDA.Abbreviations LRP late receptor potential - PDA prolonged depolarizing afterpotential - PHA prolonged hyperpolarizing afterpotential     

Design of a functional calcium channel protein: inferences about an ion channel-forming motif derived from the primary structure of voltage-gated calcium channels.

To identify sequence-specific motifs associated with the formation of an ionic pore, we systematically evaluated the channel-forming activity of synthetic peptides with sequence of predicted transmembrane segments of the voltage-gated calcium channel. The amino acid sequence of voltage-gated, dihydropyridine (DHP)-sensitive calcium channels suggests the presence in each of four homologous repeats (I-IV) of six segments (S1-S6) predicted to form membrane-spanning, alpha-helical structures. Only peptides representing amphipathic segments S2 or S3 form channels in lipid bilayers. To generate a functional calcium channel based on a four-helix bundle motif, four-helix bundle proteins representing IVS2 (T4CaIVS2) or IVS3 (T4CaIVS3) were synthesized. Both proteins form cation-selective channels, but with distinct characteristics: the single-channel conductance in 50 mM BaCl2 is 3 pS and 10 pS. For T4CaIVS3, the conductance saturates with increasing concentration of divalent cation. The dissociation constants for Ba2+, Ca2+, and Sr2+ are 13.6 mM, 17.7 mM, and 15.0 mM, respectively. The conductance of T4CaIVS2 does not saturate up to 150 mM salt. Whereas T4CaIVS3 is blocked by microM Ca2+ and Cd2+, T4CaIVS2 is not blocked by divalent cations. Only T4CaIVS3 is modulated by enantiomers of the DHP derivative BayK 8644, demonstrating sequence requirement for specific drug action. Thus, only T4CaIVS3 exhibits pore properties characteristic also of authentic calcium channels. The designed functional calcium channel may provide insights into fundamental mechanisms of ionic permeation and drug action, information that may in turn further our understanding of molecular determinants underlying authentic pore structures.