ALBERT

Notes: Seasonal variation in δ13C and δ18O of cellulose (δ13Cc and δ18Oc) was measured within two annual rings of Pinus radiata growing at three sites in New Zealand. In general, both δ13Cc and δ18Oc increased to a peak over summer. The three sites differed markedly in annual water balance, and these differences were reflected in δ13Cc and δ18Oc. Average δ13Cc and δ18Oc from each site were positively related, so that the driest site had the most enriched cellulose. δ13Cc and δ18Oc were also related within each site, although both the slope and the closeness of fit of the relationship varied between sites. Supporting the theory, the site with the lowest average relative humidity also had the greatest change in δ18Oc‰ change in δ13Cc. Specific climatic events, such as drought or high rainfall, were recorded as a peak or a trough in enrichment, respectively. These results suggest that seasonal and between-site variation in δ13Cc and δ18Oc are driven by the interaction between variation in climatic conditions and soil water availability, and plant response to this variation.

Notes: A mathematical model of stomatal conductance is presented. It is based on whole-plant and epidermal hydromechanics, and on two hypotheses: (1) the osmotic gradient across guard cell membranes is proportional to the concentration of ATP in the guard cells; and (2) the osmotic gradient that can be sustained per unit of ATP is proportional to the turgor pressure of adjacent epidermal cells. In the present study, guard cell [ATP] is calculated using a previously published model that is based on a widely used biochemical model of C3 mesophyll photosynthesis. The conductance model for Vicia faba L. is parameterized and tested As with most other stomatal models, the present model correctly predicts the stomatal responses to variations in transpiration rate, irradiance and intercellular CO2. Unlike most other models, however, this model can predict the transient stomatal opening often observed before conductance declines in response to decreases in humidity, soil water potential, or xylem conductance. The model also explicitly accommodates the mechanical advantage of the epidermis and correctly predicts that stomata are relatively insensitive to the ambient partial pressure of oxygen, as a result of the assumed dependence on ATP concentration.

Notes: In this paper, we present an integrated account of the diurnal variation in the stable isotopes of water (δD and δ18O) and dry matter (δ15N, δ13C, and δ18O) in the long-distance transport fluids (xylem sap and phloem sap), leaves, pod walls, and seeds of Lupinus angustifolius under field conditions in Western Australia. The δD and δ18O of leaf water showed a pronounced diurnal variation, ranging from early morning minima near 0‰ for both δD and δ18O to early afternoon maxima of 62 and 23‰, respectively. Xylem sap water showed no diurnal variation in isotopic composition and had mean values of −13·2 and −2·3‰ for δD and δ18O. Phloem sap water collected from pod tips was intermediate in isotopic composition between xylem sap and leaf water and exhibited only a moderate diurnal fluctuation. Isotopic compositions of pod wall and seed water were intermediate between those of phloem and xylem sap water. A model of average leaf water enrichment in the steady state (Craig & Gordon, pp. 9–130 in Proceedings of a Conference on Stable Isotopes in Oceanographic Studies and Palaeotemperatures, Lischi and Figli, Pisa, Italy, 1965; Dongmann et al., Radiation and Environmental Biophysics 11, 41–52, 1974; Farquhar & Lloyd, pp. 47–70 in Stable Isotopes and Plant Carbon–Water Relations, Academic Press, San Diego, CA, USA, 1993) agreed closely with observed leaf water enrichment in the morning and early afternoon, but poorly during the night. A modified model taking into account non-steady-state effects (Farquhar and Cernusak, unpublished) gave better predictions of observed leaf water enrichments over a full diurnal cycle. The δ15N, δ13C, and δ18O of dry matter varied appreciably among components. Dry matter δ15N was highest in xylem sap and lowest in leaves, whereas dry matter δ13C was lowest in leaves and highest in phloem sap and seeds, and dry matter δ18O was lowest in leaves and highest in pod walls. Phloem sap, leaf, and fruit dry matter δ18O varied diurnally, as did phloem sap dry matter δ13C. These results demonstrate the importance of considering the non-steady-state when modelling biological fractionation of stable isotopes in the natural environment.

Notes: Gas exchange experiments were performed with 13 plant species that differ from each other in growth-form and natural habitat. These comprised three herbaceous species, two ferns, two temperate deciduous trees, five rainforest trees and one liana from wet tropical forest. The aims were to investigate whether plants of similar growth-form and from similar habitats tended to respond similarly to a change in leaf-to-air vapour pressure difference (VPD), and to compare their ratio of intercellular to ambient partial pressures of CO2 for given conditions. Leaves were subjected to a step change in VPD and the initial and final steady rates of transpiration were used to calculate an index of sensitivity, φ, which enabled comparison of species. The results suggest that species of similar growth-form and habitat respond similarly to increasing VPD, with the temperate deciduous trees undergoing a greater reduction in stomatal conductance than the herbaceous plants in well-watered soil. Also, for these experimental conditions, the ratio of leaf internal to ambient CO2 partial pressure (pi/pa) was positively correlated with both CO2 assimilation rate and stomatal insensitivity to VPD, across the 13 species. The results are discussed in terms of growth strategies and possible advantages and limitations of hydraulic systems in different plants.

Notes: The relationship between stomatal aperture (a) and guard cell pressure (Pg) was measured directly in four different species (Vicia faba, Tradescantia virginiana, Ginkgo biloba and Nephrolepis exaltata) using a special cell pressure probe technique. The effect of epidermal turgor (Pep) on this relationship was also measured in T. virginiana. The relationship was sigmoidal for V. faba and T. virginiana, but entirely convex for G. biloba and N. exaltata. Epidermal turgor was found to have a pronounced closing effect on stomata of T. virginiana. Maximum aperture with full epidermal turgor (0·92 MPa) was about half that with zero epidermal turgor. Also, with full epidermal turgor stomata of T. virginiana did not begin to open until Pg was more than 1·25 MPa. These characteristics were used to develop an expression for a as a function of Pg and Pep. Results for the different species are compared and discussed in terms of possible advantages and limitations of water economy.

Notes: Pressure within guard cells in strips of intact epidermis of Tradescantia virginiana was controlled with a pressure probe apparatus after the guard cells had been filled with silicone oil. Pressure was increased and decreased incrementally between 0.0 and 4.1 MPa to cause inflation and deflation of the guard cells. At steady-state guard cell pressures, the width of the stomatal pore was recorded and plotted against pressure. The pressure required for near-maximum aperture was 4.1 MPa. Aperture as a function of pressure was sigmoidal.

Notes: Abstract. Analysis is made of leaf numbers observed in the field for several wheat genotypes over a wide range of sowing dates. Within any treatment, leaf number was a linear function of thermal time from seedling emergence (°C.d, the mean daily temperatures summed above a base temperature of 0°C) but the slope of the relation depended on sowing date. Leaf appearance rate doubled from October to February sowings, for which it was maximum, and usually decreased for later sowings. However, when expressed on the basis of photothermal time (°C.dl, temperatures accumulated during the light time period only), much of the sowing date effect was removed, except for the latest spring sowings. For these late sowings, leaf appearance rate was usually slower than expected. The authors conclude that both temperature and photoperiod are determinants of leaf appearance rate, and suggest that the slow rate for late sowings may be caused by a non-linear response to the higher temperatures experienced by the plant during that part of the year. The genotypic differences indicate that it may also be associated with a lack of vernalization. The use of photothermal time rather than thermal time significantly improves prediction of leaf number. However, it still provides only an empirical description of the dependence of leaf number on time and climate.

Notes: A model has been derived for the enrichment of heavy isotopes of water in leaves, including progressive enrichment along the leaf. In the model, lighter water is preferentially transpired leaving heavier water to diffuse back into the xylem and be carried further along the leaf. For this pattern to be pronounced, the ratio of advection to diffusion (Péclet number) has to be large in the longitudinal direction, and small in the radial direction. The progressive enrichment along the xylem is less than that occurring at the sites of evaporation in the mesophyll, depending on the isolation afforded by the radial Péclet number. There is an upper bound on enrichment, and effects of ground tissue associated with major veins are included. When transpiration rate is spatially nonuniform, averaging of enrichment occurs more naturally with transpiration weighting than with area-based weighting. This gives zero average enrichment of transpired water, the modified Craig–Gordon equation for average enrichment at the sites of evaporation and the Farquhar and Lloyd (In Stable Isotopes and Plant Carbon-Water Relations, pp. 47–70. Academic Press, New York, USA, 1993) prediction for mesophyll water. Earlier results on the isotopic composition of evolved oxygen and of retro-diffused carbon dioxide are preserved if these processes vary in parallel with transpiration rate. Parallel variation should be indicated approximately by uniform carbon isotope discrimination across the leaf.