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Evolutionarily stable leaf area production in plant populations.

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.

Biological Evolution↗

Growth and morphogenesis at the vegetative shoot apex of Anagallis arvensis L.

A non-destructive replica method and a 3-D reconstruction algorithm are used to analyse the geometry and expansion of the shoot apex surface. Surface expansion in the central zone of the apex is slow and nearly isotropic while surface expansion in the peripheral zone is more intense and more anisotropic. Within the peripheral zone, the expansion rate, expansion anisotropy, and the direction of maximal expansion vary according to the age of adjacent leaf primordia. For each plastochron, this pattern of expansion is rotated around the apex by the Fibonacci angle. Early leaf primordium development is divided into four stages: bulging, lateral expansion, separation, and bending. These stages differ in their geometry and expansion pattern. At the bulging stage, the site of primordium initiation shows an intensified expansion that is nearly isotropic. The following stages develop sharp meridional gradients of expansion rates and anisotropy. The adaxial primordium boundary inferred from the surface curvature is shifting until the separation stage, when a crease develops between the primordium and the apex dome. The cells forming the crease, i.e. the future leaf axil, expand along the axil and contract across it. Thus they are arrested in this unique position.

Algorithms↗

Acclimation response of spring wheat in a free-air CO(2) enrichment (FACE) atmosphere with variable soil nitrogen regimes. 3. Canopy architecture and gas exchange.

The response of whole-canopy net CO(2) exchange rate (CER) and canopy architecture to CO(2) enrichment and N stress during 1996 and 1997 for open-field-grown wheat ecosystem (Triticum aestivum L. cv. Yecora Rojo) are described. Every Control (C) and FACE (F) CO(2) treatment (defined as ambient and ambient +200 mumol mol(-1), respectively) contained a Low- and High-N treatment. Low-N treatments constituted initial soil content amended with supplemental nitrogen applied at a rate of 70 kg N ha(-1) (1996) and 15 kg N ha(-1) (1997), whereas High-N treatments were supplemented with 350 kg N ha(-1) (1996 and 1997). Elevated CO(2) enhanced season-long carbon accumulation by 8% and 16% under Low-N and High-N, respectively. N-stress reduced season-long carbon accumulation 14% under ambient CO(2), but by as much as 22% under CO(2) enrichment. Averaging both years, green plant area index (GPAI) peaked approximately 76 days after planting at 7.13 for FH, 6.00 for CH, 3.89 for FL, and 3.89 for CL treatments. Leaf tip angle distribution (LTA) indicated that Low-N canopies were more erectophile than those of High-N canopies: 48 degrees for FH, 52 degrees for CH, and 58 degrees for both FL and CL treatments. Temporal trends in canopy greenness indicated a decrease in leaf chlorophyll content from the flag to flag-2 leaves of 25% for FH, 28% for CH, 17% for CL, and 33% for FL during 1997. These results indicate that significant modifications of canopy architecture occurs in response to both CO(2) and N-stress. Optimization of canopy architecture may serve as a mechanism to diminish CO(2) and N-stress effects on CER.

Journal Article↗

Effective penumbra and scalloping effect: a dosimetric study in multifield radiotherapy with multileaf collimator for prostate cancer treatment.

AIMS AND BACKGROUND: The aim of the study was to test whether the multileaf collimator can be used to replace divergent alloy blocks for field shaping and to evaluate the dosimetric impact of the scalloping effect and the effective penumbra in multifield isocentric configurations routinely used at our department to treat early stage prostate tumor in supine patients. Deviations between measurements and treatment planning system calculations are also discussed in order to estimate the accuracy of effective penumbra calculations carried out by the software in the presence of blocks or a multileaf collimator. METHODS: The multileaf collimator installed on a dual energy (6 and 18 MV) linear accelerator Varian CLINAC 2100 C/D is an add-on component positioned below the standard jaws, with 40 computer-controlled opposed pairs of tungsten leaves. Transmission, effective penumbra and scalloping measurements were performed with films placed at different depths in a (30 x 30 x 20 cm3) acrylic phantom. A laser scanning photodensitometer was used to obtain the optical density and the relative dose profile. Effective penumbra and scalloping effect measured data were tested on a software phantom; the phantom, automatically performed with the treatment planning system, was a regular parallelepiped measuring 30 x 30 x 20 cm3 and the acrylic electronic density value. RESULTS: For one multileaf collimator-shaped field, the width of effective penumbra was about 2 mm wider than penumbra for cerrobend blocks, at a 45 degrees angle between leaf motion direction and the field edge. Collimator rotation, automatically performed by the treatment planning system, by minimizing the sum of over- and underblocked areas, reduces the differences between the multileaf collimator and blocks. The differences between measured and treatment planning system calculated data were within the treatment planning system dose calculation accuracy limits, as recommended in ICRU Report No. 42. CONCLUSIONS: Penumbra and, for the multileaf collimator, scalloping effect values seem to depend on the gantry angle, i.e., on the fields path of entry. The values for even fields are higher than the those for odd fields, because the dose gradient at the target edges is steeper for a single field than for two opposite fields, and the interplay of doses from the individual beams increases the distance between the isodose levels. Therefore, in order to reduce scalloping and effective penumbra values, it would be better to rotate the collimators, above all in even-number field techniques. In particular, the six-field technique, used mostly in prostate treatment, shows the same effective penumbra values with the multileaf collimator, with proper collimator rotation, and in blocked fields.

Humans↗

Preclinical evaluation of the reliability of a 50 MeV racetrack microtron.

PURPOSE: A 50 MeV racetrack microtron has been installed and tested at Memorial Sloan-Kettering Cancer Center. It is designed to execute multi-segment conformal therapy automatically under computer control using scanned X ray and electron beams from 10 to 50 MeV. Prior to acceptance of the machine from the manufacturer, formal reliability testing was carried out. Only in this way could confidence be gained in its usefulness for routine 3D computer-controlled conformal therapy. MATERIALS AND METHODS: To assess reliability, a set of 25 multi-segment test cases, each consisting of 10 to 17 fixed segments, was developed. The field arrangements and modalities for some of the test cases were identical to 3D conformal treatments that were being delivered with multiple static fields on conventional linear accelerators at our institution. Other cases were designed to explore reliability under more complex sets of conditions. These cases were "treated" repeatedly during a total period of 45 hours, over 5 days. During the treatments, ion chambers attached to the head of the machine provided dosimetric data for each field. Data from sensors connected to every set-up parameter (for example, couch positions, gantry angle, collimator leaf positions, etc.) were recorded and verified by an external computer. RESULTS: While preliminary tests indicated an interlock rate of 5%, final reliability test results demonstrated an interlock fault rate of approximately 0.5%. The reproducibility of dosimetric data and geometric setup parameters was within specifications. As an example, leaf position reproducibility in the patient plane was within 0.5 mm for 97% of the setups. The times required to carry out treatments were recorded and compared with the times to carry out identical treatments on a conventional linear accelerator with cerrobend blocks. Areas where additional time savings can be achieved were identified. CONCLUSIONS: As an integral part of acceptance testing, the Scanditronix MM50 was rigorously tested for reliability. The machine successfully passed these tests, providing increased confidence in its usefulness for routine 3D conformal therapy.

Humans↗

Photosynthesis and canopy characteristics in genetically defined families of silver birch (Betula pendula).

Net photosynthetic rates (A) of leaves in upper and lower crown layers (A(upper) and A(lower)), leaf area index (LAI), mean tilt angle (MTA), several leaf characteristics, and volume growth were observed in fast- and slow-growing families of a 14-year-old full-sib and half-sib family progeny test of Betula pendula Roth. Each measure of net photosynthetic rate was calculated after correcting measured net photosynthesis for the effects of environmental variables. The differences in A(upper) and LAI among families were significant. The proportions of the total variance assigned to family for A(upper), A(lower) and LAI were 33.64, 28.93 and 54.99%, respectively. The mean A(upper) and LAI of the fast-growing families were significantly higher than those of the slow-growing families, whereas the mean A(lower) of the fast-growing families was significantly lower than that of the slow-growing families. There were also significant differences among families in leaf size, leaf shape, and the ratios leaf fresh weight/area and leaf dry weight/area. Between 27.55 and 54.55% of the total variance in these characteristics could be assigned to the family effect. Volume growth was positively correlated with A(upper) and LAI, but it was most strongly correlated with A(upper) x LAI.

Journal Article↗

On estimating canopy photosynthesis and stomatal conductance in a deciduous forest with clumped foliage.

The foliage in a fully-leafed deciduous forest canopy is clumped. Consequently, theory indicates that the probability of beam penetration will be estimated more accurately with a model based on the negative binomial distribution than with a model based on the Poisson distribution, incorporating an assumption of a spherical leaf inclination angle distribution. Flux densities of photosynthetically active radiation (PAR) were measured in and above a deciduous forest canopy and were computed with the canopy radiative transfer models based on the negative binomial and Poisson distributions. These radiation values were used to compute canopy photosynthesis and stomatal conductance. Canopy photosynthesis and stomatal conductance, based on the negative binomial model, overestimated values computed from measured PAR profiles by 8 and 9%. respectively. The canopy photosynthesis and stomatal conductance values computed with the spherical Poisson model under-estimated measured values by 17 and 10%, respectively. Thus, the negative binomial radiative transfer model improves estimates of canopy photosynthesis and, to a lesser extent, stomatal conductance, inside a deciduous forest.

Journal Article↗

Influence of crown structural properties on PAR absorption, photosynthesis, and transpiration in Sitka spruce: application of a model (MAESTRO).

The structure of a tree crown can be described by the spatial distribution, inclination, and orientation of all the phytoelements (leaves, twigs, branches, trunk, etc.), and their geometric properties. The following four structural properties have been studied in relation to radiation absorption, photosynthesis, and transpiration using a simulation model named MAESTRO: crown shape, total area of leaves and their spatial distribution within the tree crown, and the leaf inclination angle distribution. It was found that the total area of leaves and their spatial distribution within the tree crown are far more important than the other two properties for radiation absorption, photosynthesis, and transpiration.

Journal Article↗

Superhydrophobic effects of self-assembled monolayers on micropatterned surfaces: 3-D arrays mimicking the lotus leaf.

The contact angle of water has been measured on a series of self-assembled monolayers (SAM) on thermally evaporated and sputter coated silver surfaces. It is found that micropatterning the surface using nanosphere lithography leads to large increases in the contact angle and generates superhydrophobic surfaces with contact angles >150 degrees. The type of functional groups on the SAMs, the metal island size, and the metal island thickness all contribute to the measured contact angle. The maximum contact angle found was 161 degrees for a fluorinated alkanethiol on 80 nm thick silver islands.

Alkanes↗

Phase Behavior of Chloroplast and Microsomal Membranes during Leaf Senescence.

Wide angle x-ray diffraction of chloroplast and microsomal membranes from primary leaves of Phaseolus vulgaris has revealed that for both types of membrane, portions of the lipid become crystalline as the tissue senesces. For young leaves the transition temperature is about 23 C for microsomes and below -30 C for chloroplast membranes, indicating that at physiological temperature the lipid is entirely liquid-crystalline. Between 2 and 3 weeks after planting the transition temperature rises to 38 C for microsomes, but for chloroplasts this increase to a point above physiological temperature does not occur until between 3 and 4 weeks. Thereafter the transition temperature continues to rise for both types of membrane with advancing senescence, although the rate of increase is greater for chloroplasts than for microsomes. The appearance at physiological temperature of gel phase lipid in the microsomes coincides temporally with the initiation of a decline in total protein in the tissue, and the incidence of crystallinity in chloroplasts coincides with loss of chlorophyll. This change in phase behavior cannot be attributed to an alteration in fatty acid composition, but for both membrane systems it correlates with an increase of about 4-fold in the sterol to phospholipid ratio.

Journal Article↗

Three-dimensional lamina architecture alters light-harvesting efficiency in Fagus: a leaf-scale analysis.

Modification of foliage exposition and morphology by seasonal average integrated quantum flux density (Qint) was investigated in the canopies of the shade-tolerant late-successional deciduous tree species Fagus orientalis Lipsky and Fagus sylvatica L. Because the leaves were not entirely flat anywhere in the canopy, the leaf lamina was considered to be three-dimensional and characterized by the cross-sectional angle between the leaf halves (theta). Both branch and lamina inclination angles with respect to the horizontal scaled positively with irradiance in the canopy, allowing light to penetrate to deeper canopy horizons. Lamina cross-sectional angle varied from 170 degrees in the most shaded leaves to 90-100 degrees in leaves in the top of the canopy. Thus, the degree of leaf rolling increased with increasing Qint, further reducing the light-interception efficiency of the upper-canopy leaves. Simulations of the dependence of foliage light-interception efficiency on theta demonstrated that decreases in theta primarily reduce the interception efficiency of direct irradiance, but that diffuse irradiance was equally efficiently intercepted over the entire range of theta values in our study. Despite strong alteration in foliage light-harvesting capacity within the canopy and greater transmittance of the upper crown compared with the lower canopy, mean incident irradiances varied more than 20-fold within the canopy, indicating inherent limitations in light partitioning within the canopy. This extensive canopy light gradient was paralleled by plastic changes in foliar structure and chemistry. Leaf dry mass per unit area varied 3-4-fold between the canopy top and bottom, providing an important means of scaling foliage nitrogen contents and photosynthetic capacity per unit area with Qint. Although leaf structure versus light relationships were qualitatively similar in all cases, there were important tree-to-tree and species-to-species variations, as well as evidence of differences in investments in structural compounds within the leaf lamina, possibly in response to contrasting leaf water availability in different trees.

Fagus↗

Regional variation in surface properties of Norway spruce and scots pine needles in relation to forest decline.

Measurements of leaf wettability (contact angle), amounts of epicuticular wax and of surface dust are reported for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L) Karst) trees growing at 12 sites in Europe from SW Germany to NE Scotland. At each site, three year classes (current year, 1 and 2-year-old) of needles were sampled from the mid-crown of up to 12 trees. Trees were selected at random from two strata, those showing visible decline symptoms (i.e. loss of needles or discoloration) and those apparently healthy. Needles for analysis were taken from apparently healthy (green) shoots in both cases. There were no significant differences between 'declining' and 'healthy' trees within sites, suggesting that leaf surface properties reflect environmental exposure rather than plant response. There were significant differences between sites, particularly for Norway spruce, which may be related to environmental factors including air pollution. Contact angles were consistently smaller at low-altitude sites in Britain and The Netherlands than at high-altitude sites in Germany where forests show decline symptoms. Leaf wettability decreased (contact angles increased) with wax amount and increased with dust amount. Leaf surface properties integrate environmental influences over long periods, and may be useful in identifying sites 'at risk' of developing decline symptoms, but causal relationships cannot be deduced without further direct experimentation.

Journal Article↗

Abutment region dosimetry for serial tomotherapy.

PURPOSE: A commercial intensity modulated radiation therapy system (Corvus, NOMOS Corp.) is presently used in our clinic to generate optimized dose distributions delivered using a proprietary dynamic multileaf collimator (DMLC) (MIMiC) composed of 20 opposed leaf pairs. On our accelerator (Clinac 600C/D, Varian Associates, Inc.) each MIMiC leaf projects to either 1.00 x 0.84 or 1.00 x 1.70 cm2 (depending on the treatment plan and termed 1 cm or 2 cm mode, respectively). The MIMiC is used to deliver serial (axial) tomotherapy treatment plans, in which the beam is delivered to a nearly cylindrical volume as the DMLC is rotated about the patient. For longer targets, the patient is moved (indexed) between treatments a distance corresponding to the projected leaf width. The treatment relies on precise indexing and a method was developed to measure the precision of indexing devices. A treatment planning study of the dosimetric effects of incorrect patient indexing and concluded that a dose heterogeneity of 10% mm(-1) resulted. Because the results may be sensitive to the dose model accuracy, we conducted a measurement-based investigation of the consequences of incorrect indexing using our accelerator. Although the indexing provides an accurate field abutment along the isocenter, due to beam divergence, hot and cold spots will be produced below and above isocenter, respectively, when less than 300 degree arcs were used. A preliminary study recently determined that for a 290 degree rotation in 1 cm mode, 15% cold and 7% hot spots were delivered to 7 cm above and below isocenter, respectively. This study completes the earlier work by investigating the dose heterogeneity as a function of position relative to the axis of rotation, arc length, and leaf width. The influence of random daily patient positioning errors is also investigated. METHODS AND MATERIALS: Treatment plans were generated using 8.0 cm diameter cylindrical target volumes within a homogeneous rectilinear film phantom. The plans included both 1 and 2 cm mode, optimized for 300 degrees, 240 degrees, and 180 degrees gantry rotations. Coronal-oriented films were irradiated throughout the target volumes and scanned using a laser film digitizer. The central target irradiated in 1 cm mode was also used to investigate the effects of incorrect couch indexing. RESULTS: The dose error as a function of couch index error was 25% mm(-1), significantly greater than previously reported. The clinically provided indexing system yielded 0.10 mm indexing precision. The intrinsic dose distributions indicated that more heterogeneous dose distributions resulted from the use of smaller gantry angle ranges and larger leaf projections. Using 300 degrees gantry angle and 1 cm mode yielded 7% hot and 15% cold spots 7 cm below and above isocenter, respectively. When a 180 degree gantry angle was used, the values changed to 22% hot and 27% cold spots for the same locations. The heterogeneities for the 2 cm mode were 70% greater than the corresponding 1 cm values. CONCLUSIONS: While serial tomotherapy is used to deliver highly conformal dose distributions, significant dosimetric factors must be considered before treatment. The patient must be immobilized during treatment to avoid dose heterogeneities caused by incorrect indexing due to patient movement. Even under ideal conditions, beam divergence can cause significant abutment-region dose heterogeneities. The use of larger gantry angle ranges, smaller leaf widths, and appropriate locations of the gantry rotation axis can minimize these effects.

Equipment Design↗

Penetration of stomata by liquids: dependence on surface tension, wettability, and stomatal morphology.

Wettability of the leaf surface, surface tension of the liquid, and stomatal morphology control penetration of stomata by liquids. The critical surface tension of the lower leaf surface of Zebrina purpusii Brückn. was estimated to be 25 to 30 dyne cm(-1). Liquids having a surface tension less than 30 dyne cm(-1) gave zero contact angle on the leaf surface and infiltrated stomata spontaneously while liquids having a surface tension greater than 30 dyne cm(-1) did not wet the leaf surface and failed to infiltrate stomata. Considering stomata as conical capillaries, we were able to show that with liquids giving a finite contact angle, infiltration depended solely on the relationship between the magnitude of the contact angle and the wall angle of the aperture. Generally, spontaneous infiltration of stomata will take place when the contact angle is smaller than the wall angle of the aperture wall. The degree of stomatal opening (4, 6, 8, or 10 mum) was of little importance. Cuticular ledges present at the entrance to the outer vestibule and between the inner vestibule and substomatal chamber resulted in very small if not zero wall angles, and thus played a major role in excluding water from the intercellular space of leaves. We show why the degree of stomatal opening cannot be assessed by observing spontaneous infiltration of stomata by organic liquids of low surface tension.

Journal Article↗

Reflectance and transmittance of light by leaves.

Spectrophotometric transmittance and reflectance curves were recorded for wavelengths from 0.45 (in some cases 0.34) to 2.7 micrometers for faces and backs of leaves and for stacked leaves of several plant species. Measurements were made at different angles of illumination. Leaf spectrophotometric curves were compared with curves for leaf extracts, potato tuber tissue, glass beads in water, and frozen leaves to demonstrate the physical bases for the leaf curves. Leaves were infiltrated with liquids of different refractive indices for further comparison of spectrophotometric curves. Goniophotometric reflectance curves were recorded, giving visible reflectance and degree of polarization as functions of viewing angle for two different angles of illumination.No retroreflection was observed, and no phenomena were observed which could be attributed to interference because of similarity between leaf structural sizes and wavelengths used.

Journal Article↗

Surface characterization and adhesion and friction properties of hydrophobic leaf surfaces.

Super-hydrophobic surfaces as well as low adhesion and friction are desirable for various industrial applications. Certain plant leaves are known to be hydrophobic in nature. These leaves are hydrophobic due to the presence of microbumps and a thin wax film on the surface of the leaf. The purpose of this study is to fully characterize the leaf surface and to separate out the effects of the microbumps and the wax on the hydrophobicity. Furthermore, the adhesion and friction properties of the leaves, with and without wax, are studied. Using an optical profiler and an atomic/friction force microscope (AFM/FFM), measurements on the hydrophobic leaves, both with and without wax, were made to fully characterize the leaf surface. Using a model that predicts contact angle as a function of roughness, the roughness factor for the hydrophobic leaves has been calculated, which is used to calculate the contact angle for a flat leaf surface. It is shown that both the microbumps and the wax play an equally important role in the hydrophobic nature as well as adhesion and friction of the leaf. This study will be useful in developing super-hydrophobic surfaces.

Adhesiveness↗

Geometric leaf placement strategies.

Geometric leaf placement strategies for multileaf collimators (MLCs) typically involve the expansion of the beam's-eye-view contour of a target by a uniform MLC margin, followed by movement of the leaves until some point on each leaf end touches the expanded contour. Film-based dose-distribution measurements have been made to determine appropriate MLC margins--characterized through an index d90--for multileaves set using one particular strategy to straight lines lying at various angles to the direction of leaf travel. Simple trigonometric relationships exist between different geometric leaf placement strategies and are used to generalize the results of the film work into d90 values for several different strategies. Measured d90 values vary both with angle and leaf placement strategy. A model has been derived that explains and describes quite well the observed variations of d90 with angle. The d90 angular variations of the strategies studied differ substantially, and geometric and dosimetric reasoning suggests that the best strategy is the one with the least angular variation. Using this criterion, the best straightforwardly implementable strategy studied is a 'touch circle' approach for which semicircles are imagined to be inscribed within leaf ends, the leaves being moved until the semicircles just touch the expanded target outline.

Film Dosimetry↗

Some plant leaves have orientation-dependent EPR and NMR spectra.

Proton nuclear magnetic resonance ((1)H NMR) spectra of leaves from 50 plant species were obtained at a spectrometer frequency of 470 MHz. Water present in leaf samples gives rise to characteristic spectral patterns. Most species show only one broad (1)H NMR peak; however, the leaves of some plants display complex, orientation-dependent spectra in which a common three-line pattern is discerned. The pattern varies with the angle between the leaf surface and the external magnetic field. Proton relaxation measurements show the presence of at least two water compartments in the leaves. The compartments are responsible for different components of the spectral pattern. EPR spectra, obtained at 35 GHz and at a temperature of -180 degrees C, of plant leaf sections are dominated by the strong signals of manganous ions. We find that most plant leaves have isotropic Mn(2+) EPR spectra. However, in some species (including ones that exhibit orientation-dependent (1)H NMR spectra) we detect orientation-dependent intensities in the forbidden lines; the spectra indicate that Mn(2+) ions occupy binding sites with axial or lower symmetry on nonrandomly oriented membranes. Both the NMR and the EPR results suggest that the chloroplasts of some plants are preferentially aligned with respect to the leaf surface.

Journal Article↗