ALBERT

Notes: The internal conductance from intercellular spaces to the sites of carboxylation (gi) has only been measured in a few tree species and not in conifers, despite the fact it may impose a large limitation on photosynthesis. The present study provides the first estimates of gi for a coniferous species, and examines variation in gi with height and its relationships to anatomical, biochemical and physiological traits. Measurements were made on upper and lower canopy current-year needles of 50-year-old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco). Needle thickness and specific leaf area decreased by 30% from the top to bottom of the canopy. These anatomical/morphological changes were accompanied by modest variation in allocation of N to chlorophyll and the chlorophyll a/b ratio. Allocation of N to Rubisco did not vary with height, but the ratio of Rubisco to chlorophyll did owing to the aforementioned changes in allocation to chlorophyll. The value of gi was estimated in one tree from concurrent measurements of carbon isotope discrimination and net photosynthesis. To examine the variation in gi among trees a second independent method based on day respiration and the difference between the chloroplastic and intercellular photocompensation points (photocompensation point method) was used. Estimates of gi obtained by the two methods agreed well with values varying between 0.14 and 0.20 mol m−2 s−1. It is estimated that gi limits photosynthesis by approximately 20% as compared to an approximately 30% stomatal limitation (under well-watered conditions). The value of gi scaled approximately with maximum rates of photosynthesis, which were significantly greater in upper canopy needles. Nevertheless, gi did not vary significantly with canopy height, owing to greater variability in gi than photosynthesis.

Notes: 
 A, assimilation rate
a, fractionation against 13C for CO2 diffusion through air
b, net fractionation against 13C during CO2 fixation
Ca, ambient CO2 concentration
Cc, CO2 concentration at the chloroplast
Ci, intercellular CO2 concentration
D, vapour pressure deficit
En, needle transpiration rate
Ep, whole plant water use
gw, leaf internal transfer conductance to CO2
gs, stomatal conductance to water vapour
L, projected leaf area
NUE, nitrogen use efficiency
PEP, phosphoenolpyruvate
Rubisco, ribulose-1,5-biphosphate carboxylase
TDR, time domain reflectometry
WUE, water use efficiency
Δ, carbon isotope discrimination
δ13C, carbon isotope abundance parameter
δ13Ca, carbon isotopic composition of atmospheric CO2
θ, volumetric soil water content

The effect of nitrogen stress on needle δ13C, water-use efficiency (WUE) and biomass production in irrigated and dry land white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings, representing 10 controlled crosses, were planted in the field in individual buried sand-filled cylinders. Two nitrogen treatments were imposed, nitrogen stressed and fertilized. The ranking of δ13C of the crosses was maintained across all combinations of water and nitrogen treatments and there was not a significant genetic versus environmental interaction. The positive relationships between needle δ13C, WUE and dry matter production demonstrate that it should be possible to use δ13C as a surrogate for WUE, and to select for increased WUE without compromising yield, even in nitrogen deficient environments. Nitrogen stressed seedlings had the lowest needle δ13C in both irrigated and dry land conditions. There was a positive correlation between needle nitrogen content and δ13C that was likely associated with increased photosynthetic capacity. There was some indication that decreased nitrogen supply led to increased stomatal conductance and hence lower WUE. There was a negative correlation between intrinsic water use efficiency and photosynthetic nitrogen use efficiency (NUE). This suggests that white spruce seedlings have the ability to maximize NUE when water becomes limited. There was significant genetic variation in NUE that was maintained across treatments. Our results suggest that in white spruce, there is no detectable effect of anaplerotic carbon fixation and that it is more appropriate to use a value of 29‰ (‘Rubisco only’) for the net discrimination against 13C during CO2 fixation. This leads to excellent correspondence between values of Ci/Ca derived from gas exchange measurements or from δ13C.

Notes: A simple and inexpensive feedback control system that provides continuous and precise control of photosynthetic photon flux density (PPFD) in a whole plant cuvette is described. A ‘Plexiglass’ tank is interposed between a light source and cuvette and PPFD changed by varying the level of dyed liquid in the tank. The amount of liquid pumped into or drained from the tank is a function of the difference (error) between a defined set point value of PPFD and that measured in the cuvette. The set point can be varied as a function of time, can follow the output of a quantum sensor measuring ambient PPFD or can be driven by values of PPFD read from a data file. Within the 0.4 to 0.64 μm waveband, the dye acts as a neutral density filter so that there is no change in spectral distribution with PPFD. Photosynthetic photon flux density in the cuvette was controlled to better than 20 μmol m−2s−1 when the set point was varied from 200 to 1100 μmol m−2s−1 over 3 min. When the set point was held constant or changed less rapidly, errors did not exceed 5 μmol m−2s−1. Net photosynthesis of Western redcedar (Thuja plicata Donn.) seedlings held at 18 °C closely followed rapid changes in PPFD.

Notes: Sudden but transient changes in the fraction or illuminated foliage area in a well-watered 7-year-old Pinus radiata D. Don tree were imposed by completely covering either the upper 22% or the lower 78% of the foliage for periods of up to 36 h. Measurements of transpiration flux density (E), tree conductance (gt), stomatal conductance (gs) and net photosynthesis (A) were made to test the hypothesis that compensatory responses would occur in the remaining illuminated foliage when the cover was installed. When the lower foliage was covered there was an immediate decrease in gt. However, when tree conductance was normalized with respect to the illuminated leaf area (gt'), it increased between 50 and 75%, depending on the value of air saturation deficit (D). The effect was also apparent from concurrent measurements of increases in gs and A up to 59 and 24%, respectively, for needles in the top third of (he crown. When the cover was removed these effects were reversed. The changes in the lower foliage when the upper foliage was covered were much smaller. Changes in bulk needle water potential were small. It is suggested that the observed responses occurred because of a perturbation to the hydraulic pathway in the xylem that could have triggered the action of a chemical signal to regulate stomatal conductance and photosynthesis.

Notes: Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (gc). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in gc with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.

Notes: Virtually all current estimates of the maximum carboxylation rate (Vcmax) of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and the maximum electron transport rate (Jmax) for C3 species implicitly assume an infinite CO2 transfer conductance (gi). And yet, most measurements in perennial plant species or in ageing or stressed leaves show that gi imposes a significant limitation on photosynthesis. Herein, we demonstrate that many current parameterizations of the photosynthesis model of Farquhar, von Caemmerer & Berry (Planta 149, 78–90, 1980) based on the leaf intercellular CO2 concentration (Ci) are incorrect for leaves where gi limits photosynthesis. We show how conventional A–Ci curve (net CO2 assimilation rate of a leaf –An– as a function of Ci) fitting methods which rely on a rectangular hyperbola model under the assumption of infinite gi can significantly underestimate Vcmax for such leaves. Alternative parameterizations of the conventional method based on a single, apparent Michaelis–Menten constant for CO2 evaluated at Ci[Km(CO2)i] used for all C3 plants are also not acceptable since the relationship between Vcmax and gi is not conserved among species. We present an alternative A–Ci curve fitting method that accounts for gi through a non-rectangular hyperbola version of the model of Farquhar et al. (1980). Simulated and real examples are used to demonstrate how this new approach eliminates the errors of the conventional A–Ci curve fitting method and provides Vcmax estimates that are virtually insensitive to gi. Finally, we show how the new A–Ci curve fitting method can be used to estimate the value of the kinetic constants of Rubisco in vivo is presented

Notes: The relationship among water use efficiency (WUE), productivity and carbon isotopic composition (δ13C) in white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings representing 10 controlled crosses were planted in the field in individual buried sand-filled cylinders. The soil water content in the cylinders was measured using time domain reflectometry over two growing seasons and seedling water use determined by water balance. Two watering treatments were imposed: irrigation and dry land. There was significant (1.6–2.0%c) genetic variation in needle δ13C. Ranking of crosses in terms of δ13C was generally maintained over watering treatments and there was not a significant genetic versus environmental interaction. There was a positive correlation between δ13C and both intrinsic and long-term WUE (more positive δ13C with increased WUE) and between δ13C and productivity, suggesting a correlation due to variation in photosynthetic capacity. Root to shoot ratios did not increase in water-stressed plants, indicating that responses to drought were primarily at the level of gas exchange, rather than through morphological changes. Our results indicate that it should be possible to use δ13C as a surrogate for WUE and to select white spruce genotypes for high WUE without compromising yield.

Notes: We determined the stable carbon isotope composition (δ1.3C) of cellulose extracted from early and late wood in Douglas fir [Pseudotsuga menziexii (Mirb.) Franco] tree rings. Data were obtained for the period 1962 to 1981, at the start of which the trees were 20 years old. A water balance model was used to calculate daily stand transpiration and water deficit. The model incorporates site factors (soil water availability, slope and aspect) and environmental variables (solar radiation, air temperature and rainfall). There was far greater variability in late wood than in early wood δ1.3C. In wet years, late wood δ1.3C was significantly lighter (by as much as 2δ) than early wood δ1.3C but in dry years this difference was reversed. Differences between spring and summer cumulative transpiration accounted for almost 60δ of the variability in differences between early and late wood δ1.3C. We found excellent correspondence between summer cumulative transpiration and late wood δ1.3C, with estimates of transpiration accounting for up to 93% of the variability in δ1.3C. Correlations between early wood δ1.3C and spring transpiration were generally poor (r2〈0.4), but we were able to identify those exceptional years in which there had been a very dry spring. Our results indicate that, while tree ring δ1.3C correlates reasonably well with basal area increment, it is a far better indicator of inter- and intra-annual variability in water availability than radial growth.

Notes: Mustard (Sinapis alba L.), Argentine canola (Brassica napus L. cv. Westar), Polish canola (Brassica campestris L. cv. Tobin), pea (Pisum sativum L.), durum wheat (Triticum durum L. cv. Kyle) and soft wheat (Triticum aestivum L. cv. Fielder) were grown at Outlook, Saskatchewan, Canada, under irrigated and dryland conditions. Carbon isotope discrimination (Δ) and water-use efficiency (W), defined as grams of above ground dry matter produced per kilogram water used, were negatively correlated in the six field-grown crops. In irrigated plants Δ remained relatively constant (20–21‰) throughout the growing season. However, in dryland plants, Δ declined in response to the progressive depletion of stored soil water (Polish canola, 20-2-18-8‰; mustard, 19.9–18 5‰; pea, 19.9–17 2‰ durum wheat, 19.7–16.4‰; Argentine canola, 19.4–17.6‰; soft wheat, 19.0–17.4‰). Although there were genetic differences in Δ among the species, water availability was the major factor controlling Δ.

Notes: The three dimensional distribution of intercepted radiation, intercellular CO2 concentration (Ci) and late summer needle nitrogen (N) concentration were determined at the tips of all 54 branches in a 6·2-m-tall Pinus radiata D. Don tree growing in a New Zealand plantation. Measurements included above- and below-canopy irradiance, leaf stable carbon isotopic composition (δ13C) and tree canopy architecture. The radiation absorption component of the model, MAESTRO, was tested on site and then used to determine the branch tip distribution of intercepted radiation. We hypothesized that in branch tip needles: (i) the allocation of nitrogen and other nutrients would be closely associated with the distribution of intercepted radiation, reflecting carbon gain optimization theory, and (ii) Ci would predominantly reflect changes in photosynthetic rate (A) rather than stomatal conductance (gs), indicating that the increase in A for a given increase in N concentration was larger than the corresponding increase in gs. Needle nitrogen concentration was poorly related to intercepted radiation, regardless of the period over which the latter was calculated. At a given height, there was a large azimuthal variation in intercepted radiation but N concentration was remarkably uniform around the tree canopy. There was, however, a linear and positive correspondence between N concentration and δ13C and needle height above ground (r2 = 0·73 and 0·68, respectively). The very strong linear correspondence between N concentration and Ci (r2 = 0·71) was interpreted, using gas exchange measurements, as supporting our second hypothesis. Recognizing the strong apical control in P. radiata and possible effects of leaf nitrogen storage in an evergreen species, we propose that the tree leader must have constituted a very strong carbon sink throughout the growing season, and that the proximity of branch tip needles to the leader affected their photosynthetic capacity and nutrient concentration, independent of intercepted radiation. This implies an integrated internal determination of resource allocation within the tree and challenges the current convention that resources are optimally distributed according to the profile of intercepted radiation.