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Populus: arabidopsis for forestry. Do we need a model tree?

Trees are used to produce a variety of wood-based products including timber, pulp and paper. More recently, their use as a source of renewable energy has also been highlighted, as has their value for carbon mitigation within the Kyoto Protocol. Relative to food crops, the domestication of trees has only just begun; the long generation time and complex nature of juvenile and mature growth forms are contributory factors. To accelerate domestication, and to understand further some of the unique processes that occur in woody plants such as dormancy and secondary wood formation, a 'model' tree is needed. Here it is argued that Populus is rapidly becoming accepted as the 'model' woody plant and that such a 'model' tree is necessary to complement the genetic resource being developed in arabidopsis. The genus Populus (poplars, cottonwoods and aspens) contains approx. 30 species of woody plant, all found in the Northern hemisphere and exhibiting some of the fastest growth rates observed in temperate trees. Populus is fulfilling the 'model' role for a number of reasons. First, and most important, is the very recent commitment to sequence the Populus genome, a project initiated in February 2002. This will be the first woody plant to be sequenced. Other reasons include the relatively small genome size (450-550 Mbp) of Populus, the large number of molecular genetic maps and the ease of genetic transformation. Populus may also be propagated vegetatively, making mapping populations immortal and facilitating the production of large amounts of clonal material for experimentation. Hybridization occurs routinely and, in these respects, Populus has many similarities to arabidopsis. However, Populus also differs from arabidopsis in many respects, including being dioecious, which makes selfing and back-cross manipulations impossible. The long time-to-flower is also a limitation, whilst physiological and biochemical experiments are more readily conducted in Populus compared with the small-statured arabidopsis. Recent advances in the development of large expressed sequence tagged collections, microarray analysis and the free distribution of mapping pedigrees for quantitative trait loci analysis secure Populus as the ideal subject for further exploitation by a wide range of scientists including breeders, physiologists, biochemists and molecular biologists. In addition, and in contrast to other model plants, the genus Populus also has genuine commercial value as a tree for timber, plywood, pulp and paper.

Arabidopsis↗

Does canopy position affect wood specific gravity in temperate forest trees?

The radial increases in wood specific gravity known in many tree species have been interpreted as providing mechanical support in response to the stresses associated with wind loading. This interpretation leads to the hypothesis that individuals reaching the canopy should (1) be more likely to have radial increases in specific gravity and (2) exhibit greater increases than individuals in the subcanopy. Wood specific gravity was determined for three species of forest trees (Acer rubrum, Fagus grandifolia and Tsuga canadensis) growing in central Massachusetts, USA. Acer rubrum shows radial increases in specific gravity, but these increases are not more pronounced in canopy trees; the other two species show a pattern of radial decreases. The degree of radial increase or decrease is influenced by tree height and diameter. Of the dominant tree species for which we have data, A. rubrum, Betula papyrifera and Pinus strobus show radial increases in specific gravity, whereas F. grandifolia, T. canadensis and Quercus rubra show decreases. The occurrence of radial increases in B. papyrifera and P. strobus, which are often canopy emergents, suggests that it is overall adaptive strategy that is important rather than position (canopy vs. subcanopy) of any individual tree. It is suggested that radial increases in specific gravity are associated with early-successional status or characteristics and decreases with late-successional status or persistence in mature forest.

Acer↗

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India.

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.

Biodiversity↗

Time-kill studies of tea tree oils on clinical isolates.

Tea tree oil has recently emerged as an effective topical antimicrobial agent active against a wide range of organisms. Tea tree oil may have a clinical application in both the hospital and community, especially for clearance of methicillin-resistant Staphylococcus aureus (MRSA) carriage or as a hand disinfectant to prevent cross-infection with Gram-positive and Gramnegative epidemic organisms. Our study, based on the time-kill approach, determined the kill rate of tea tree oil against several multidrug-resistant organisms, including MRSA, glycopeptide-resistant enterococci, aminoglycoside-resistant klebsiellae, Pseudomonas aeruginosa and Stenotrophomonas maltophilia, and also against sensitive microorganisms. The study was performed with two chemically different tea tree oils. One was a standard oil and the other was Clone 88 extracted from a specially bred tree, which has been selected and bred for increased activity and decreased skin irritation. Our results confirm that the cloned oil had increased antimicrobial activity when compared with the standard oil. Most results indicated that the susceptibility pattern and Gram reaction of the organism did not influence the kill rate. A rapid killing time (less than 60 min) was achieved with both tea tree oils with most isolates, but MRSA was killed more slowly than other organisms.

Anti-Infective Agents, Local↗

Increased growth of young citrus trees under reduced radiation load in a semi-arid climate.

This study investigated the effects of radiation heat-load reduction by shading on the growth and development of citrus trees in a warm subtropical region. The experiment was conducted from mid-June until late October when daily maximal air temperature averaged 29.3 degrees C. Two-year-old de-fruited Murcott tangor (Citrus reticulata BlancoxCitrus sinensis (L.) Osb.) trees were grown under 30% or 60% shade tunnels, or 60% flat shade (providing midday shade only), using highly reflective aluminized nets. Non-shaded trees were used as the control. Shading reduced direct more than diffuse radiation. Daily radiation was reduced by 35% for the 30% Tunnel and 60% Flat treatments, and by 55% for the 60% Tunnel. Two days of intensive measurement showed that shading increased average sunlit leaf conductance by 44% and photosynthesis by 29%. Shading did not significantly influence root and stem dry weight growth, but it increased the increment in leaf dry weight during the three month period by an average of 28% relative to the control, while final tree height in the 30% Tunnel treatment exceeded the control by 35%. Shoot to root and shoot mass ratios increased and root mass ratio decreased due to shading because of the increase in leaf dry weight. Shading increased starch concentration in leaves while the shadiest treatment, 60% Tunnel, decreased starch concentration in the roots. Carbon isotope ratio (delta(13)C) of exposed leaves that developed under shading was significantly reduced by 1.9 per thousand in the 60% Tunnel, indicating that shading increased CO(2) concentrations at the chloroplasts (C(c)), as would be expected from increased conductance. Substomatal CO(2) concentrations, C(i), computed from leaf net CO(2) assimilation rate and conductance values, also indicate that shading increases internal CO(2) concentrations. Based on tree dry mass, tree height, and total carbohydrates fractions, the 30% Tunnel and the 60% Flat were the optimal shade treatments.

Citrus↗

Chronic ozone exposure affects leaf senescence of adult beech trees: a chlorophyll fluorescence approach.

Accelerated leaf senescence is one of the harmful effects of elevated tropospheric ozone concentrations ([O(3)]) on plants. The number of studies dealing with mature forest trees is scarce however. Therefore, five 66-year-old beech trees (Fagus sylvatica L.) have been exposed to twice-ambient (2xambient) [O(3)] levels by means of a free-air canopy O(3) exposure system. During the sixth year of exposure, the hypothesis of accelerated leaf senescence in 2xambient [O(3)] compared with ambient [O(3)] trees was tested for both sun and shade leaves. Chlorophyll (chl) fluorescence was used to assess the photosynthetic quantum yield, and chl fluorescence images were processed to compare functional leaf homogeneity and the proportion of O(3)-injured leaf area (stipples) under ambient and 2xambient [O(3)] regimes. Based on the analysis of chl fluorescence images, sun leaves of both ambient and 2xambient [O(3)] trees had apparently developed typical necrotic O(3) stipples during high O(3) episodes in summer, while accelerated senescence was only observed with sun leaves of 2xambient [O(3)] trees. This latter effect was indicated along with a faster decrease of photosynthetic quantum yield, but without evidence of changes in non-photochemical quenching. Overall, treatment effects were small and varied among trees. Therefore, compared with ambient [O(3)], the consequence of the observed O(3)-induced accelerated leaf senescence for the carbon budget is likely limited.

Acclimatization↗

Relationships between hydraulic architecture and leaf photosynthetic capacity in nitrogen-fertilized Eucalyptus grandis trees.

We compared the effects of nitrogen fertilization on shoot hydraulic architecture and leaf photosynthetic properties of Eucalyptus grandis Hill ex Maiden trees in Hawaii. It was hypothesized that water transport capacity would adjust to nutrient availability, with leaf specific hydraulic conductivity (kl) increasing in fertilized trees in coordination with higher photosynthetic capacity per unit leaf area. Trees were grown from seedlings in the field for 10 months at four rates of nitrogen (N) fertilization between 0 and 336 kg x ha(-1). Leaf water potentials, photosynthetic capacity and kl were measured before whole shoots were harvested to determine total growth, leaf area and sapwood density. Mean tree height increased from 4 to 5.3 m, stem basal area increased from 27 to 67 cm2 and total leaf area increased from 15 to 40 m2 between the lowest and highest rates of fertilizer addition. When trees were compared on the basis of leaf nitrogen per unit area (N(area)), light-saturated rates of photosynthesis on an area and mass basis and the maximum rate of electron transport all increased from 50% to more than 100% as N(area) increased from 0.8 to 2.1 g x m(-2). Branch specific hydraulic conductivity (ks) and kl increased with height in the crown. However, there was no change in branch kl or the ratio of leaf area to sapwood area of the whole shoot in response to fertilization, and ks and density of the sapwood were unrelated to leaf N(area). In contrast to photosynthesis, stomatal conductance did not respond to fertilization, leading to decreased internal carbon dioxide partial pressure (p(i)/p(a)) in fertilized plants and similar leaf water potentials in all plants. Consistent with the behavior of p(i)/p(a), carbon isotope discrimination decreased by 2% with increasing leaf N(area), supporting the conclusion that intrinsic water-use efficiency was enhanced by fertilization. Increased growth in response to fertilization involved adjustment at the leaf level rather than a change in the balance between water transport capacity and leaf area. It is proposed that, when there are changes in leaf properties without any external change in water availability or evaporative demand, leaf photosynthesis and stomatal conductance are partially constrained by the hydraulic architecture of the tree.

Eucalyptus↗

Carbon dioxide exchange in Norway spruce at the shoot, tree and ecosystem scale.

Net CO2 exchange in a 35-year-old boreal Norway spruce (Picea abies (L.) Karst.) forest in northern Sweden was measured at the shoot (NSE), tree (NTE) and ecosystem levels (NEE) by means of shoot cuvettes, whole-tree chambers and the eddy covariance technique, respectively. We compared the dynamics of gross primary production (GPP) at the three levels during the course of a single week. The diurnal dynamics of GPP at each level were estimated by subtracting half-hourly or hourly model-estimated values of total respiration (excluding light-dependent respiration) from net CO(2) exchange. The relationship between temperature and total respiration at each level was derived from nighttime measurements of NSE, NTE and NEE over the course of 1 month. There was a strong linear relationship (r2 = 0.93) between the hourly estimates of GPP at the shoot and tree levels, but the correlation between shoot- and ecosystem-level GPP was weaker (r2 = 0.69). However, the correlation between shoot- and ecosystem-level GPP was improved (r2 = 0.88) if eddy covariance measurements were restricted to periods when friction velocity was > or = 0.5 m s(-1). Daily means were less dependent on friction velocity, giving an r2 value of 0.94 between shoot- and ecosystem-level GPP. The correlation between shoot and tree levels also increased when daily means were compared (r2 = 0.98). Most of the measured variation in carbon exchange rate among the shoot, tree and ecosystem levels was the result of periodic low coupling between vegetation and the atmosphere at the ecosystem level. The results validate the use of measurements at the shoot and tree level for analyzing the contribution of different compartments to net ecosystem CO2 exchange.

Carbon Dioxide↗

Genomics and the tree physiologist.

During the last decade, a strong case has been made for viewing trees as model systems in plant biology. More recently, forest biologists have argued for the sequencing of the genome of a forest tree. Now, the United States Department of Energy has announced plans for sequencing the genome of the Populus trichocarpa clone, "Nisqually-1." Thus, forest biology is poised to enter an exciting period of scientific discovery. Embracing new technology and new research paradigms, however, is never easy. It is, therefore, timely to ask how tree physiologists will take advantage of the Populus genome data. We contend that the most attractive opportunities will arise through genome-wide research designed to: (1). examine the differences between trees and herbaceous annuals; (2). explore questions relating to the temporal and spatial scales that characterize the life histories and growth of trees; and (3). investigate cause-and-effect relationships that are intractable to conventional research methodologies. To highlight the potential applications of genomics in tree physiology, we briefly discuss each of these approaches.

Genome, Plant↗

Tree- and needle-age-dependent variations in antioxidants and photoprotective pigments in Norway spruce needles at the alpine timberline.

To cope with environmental stress, plants are equipped with antioxidative (e.g., ascorbate, glutathione and alpha-tocopherol) and photoprotective (e.g., xanthophyll cycle pigments) defense systems. We investigated the defense capacities of three tree age classes (mature, sapling and seedling) of Norway spruce (Picea abies (L.) Karst.) at a field site near the timberline. Biochemical data were expressed on both a needle dry mass and a surface area basis. Compared with current-year needles, previous-year needles contained higher mass- and area-based concentrations of chlorophylls and alpha-tocopherol, and a larger xanthophyll cycle pool that was in a more epoxidized state. Total glutathione concentration was lower, the glutathione pool was more reduced and the ascorbate pool was more oxidized in previous-year needles than in current-year needles. Needle concentrations of glutathione and alpha-tocopherol increased and chlorophyll concentration decreased with increasing tree age when expressed on a surface area basis. On a dry mass basis, these trends were reversed or nonexistent. The ascorbate pool was more reduced and the glutathione pool was more oxidized in needles of mature trees than in needles of saplings and seedlings. The proportion of protective xanthophyll cycle pigments decreased and the de-epoxidation state increased with increasing tree age. We conclude that tree age and the basis of expression of antioxidant concentration--surface area or dry mass--are important in scaling from seedlings to large trees.

Antioxidants↗

Interaction between sapwood and foliage area in alpine ash (Eucalyptus delegatensis) trees of different heights.

In native stands of Eucalyptus delegatensis R. T. Baker, sapwood area (As) to foliage area (Af) ratios (As:Af) decreased as tree height increased, contradicting the common interpretation of the Pipe Model Theory as well as the generally observed trend of increasing As:Af ratios with tree height. To clarify this relationship, we estimated sapwood hydraulic conductivity theoretically based on measurements of sapwood vessel diameters and Poiseuille's law for fluid flow through pipes. Despite the observed decrease in As:Af ratios with tree height, leaf specific conductivity increased with total tree height, largely as a result of an increase in the specific conductivity of sapwood. This observation supports the proposition that the stem's ability to supply foliage with water must increase as trees grow taller, to compensate for the increased hydraulic path length. The results presented here highlight the importance of measuring sapwood hydraulic conductivity in analyses of sapwood-foliage interactions, and suggest that measurements of sapwood hydraulic conductivity may help to resolve conflicting observations of how As:Af ratios change as trees grow taller.

Eucalyptus↗

Tracing carbon uptake from a natural CO2 spring into tree rings: an isotope approach.

We analyzed 14C, 13C and 18O isotope variations over a 50-year period in tree rings of Quercus ilex L. trees growing at a natural CO2 spring in a Mediterranean ecosystem. We compared trees from two sites, one with high and one with low exposure to CO2 from the spring. The spring CO2 is free of 14C. Thus, this carbon can be traced in the wood, and the amount originating from the spring calculated. The amount decreased over time, from about 40% in 1950 to 15% at present for the site near the spring, indicating a potential difficulty in the use of natural CO2 springs for elevated CO2 research. The reason for the decrease may be decreasing emission from the spring or changes in stand structure, e.g., growth of the canopy into regions with lower concentrations. We used the 14C-calculated CO2 concentration in the canopy to determine the 13C discrimination of the plants growing under elevated CO2 by calculating the effective canopy air 13C/12C isotopic composition. The trees near the spring showed a 2.5 per thousand larger 13C discrimination than the more distant trees at the beginning of the investigated period, i.e., for the young trees, but this difference gradually disappeared. Higher discrimination under elevated CO2 indicated reduced photosynthetic capacity or increased stomatal conductance. The latter assumption is unlikely as inferred from the 18O data, which were insensitive to CO2 concentration. In conclusion, we found evidence for a downward adjustment of photosynthesis under elevated CO2 in Q. ilex in this dry, nutrient-poor environment.

Carbon Dioxide↗

Nitrogen storage and its interaction with carbohydrates of young apple trees in response to nitrogen supply.

Bench-grafted 'Fuji/M.26' apple (Malus domestica Borkh.) trees received a constant nitrogen (N) supply (10.7 mM) from bud break to the end of June, and were then fertigated with 0, 5, 10, 15 or 20 mM N in a modified Hoagland's solution for 2 months during the summer. In mid-October, half of the trees fertigated at each N concentration were sprayed twice with 3% urea, whereas the remaining trees served as controls. All trees were harvested after natural leaf fall and were stored at 2 degrees C. Five trees from each of the N treatment combinations were destructively sampled during dormancy to determine the composition of N and total nonstructural carbohydrates (TNC). As the N supply from fertigation increased, amounts of N in both free amino acids and proteins increased, whereas C/N ratios decreased. Foliar urea applications in the fall significantly increased amounts of N in both free amino acids and proteins, but decreased their C/N ratios. Arginine, the most abundant amino acid in both free amino acids and in proteins, accounted for an increasing proportion of N in free amino acids and proteins with increasing N supply from fertigation or foliar urea application. The ratio of protein N to free amino acid N decreased from about 27.1 to 3.2 as N supply from fertigation increased from 0 to 20 mM, and decreased further to 3.0 in response to foliar urea applications in the fall. Concentrations of glucose, fructose, sucrose and TNC decreased as the N supply from fertigation increased, whereas concentrations of sorbitol and starch remained relatively unchanged. Foliar urea applications decreased the concentration of each TNC component and the TNC concentration in each N fertigation treatment. A negative linear relationship was found between carbon in TNC and N in proteins and free amino acids. The sum of carbon in TNC, proteins and free amino acids remained constant in response to N supply from fertigation. However, foliar urea applications decreased the sum of carbon in proteins, free amino acids and TNC because about 21% of the decrease in TNC carbon was not recovered in free amino acids or proteins. Young apple trees store N and carbon dynamically in response to N supply. As N supply increases, an increasing proportion of N is found in the form of free amino acids, which have a low carbon cost, although proteins remain the main form of N storage. Furthermore, part of the carbon from TNC is incorporated into amino acids and proteins, decreasing the carbon stored as TNC and increasing the carbon stored as amino acids and proteins.

Carbohydrate Metabolism↗

Laser ablation-combustion-GC-IRMS--a new method for online analysis of intra-annual variation of delta13C in tree rings.

We present a new, rapid method for high-resolution online determination of delta13C in tree rings, combining laser ablation (LA), combustion (C), gas chromatography (GC) and isotope ratio mass spectrometry (IRMS) (LA-C-GC-IRMS). Sample material was extracted every 6 min with a UV-laser from a tree core, leaving 40-microm-wide holes. Ablated wood dust was combusted to CO2 at 700 degrees C, separated from other gases on a GC column and injected into an isotope ratio mass spectrometer after removal of water vapor. The measurements were calibrated against an internal and an external standard. The tree core remained intact and could be used for subsequent dendrochronological and dendrochemical analyses. Cores from two Scots pine trees (Pinus sylvestris spp. sibirica Lebed.) from central Siberia were sampled. Inter- and intra-annual patterns of delta13C in whole-wood and lignin-extracted cores were indistinguishable apart from a constant offset, suggesting that lignin extraction is unnecessary for our method. Comparison with the conventional method (microtome slicing, elemental analysis and IRMS) indicated high accuracy of the LA-C-GC-IRMS measurements. Patterns of delta13C along three parallel ablation lines on the same core showed high congruence. A conservative estimate of the precision was +/- 0.24 per thousand. Isotopic patterns of the two Scots pine trees were broadly similar, indicating a signal related to the forest stand's climate history. The maximum variation in delta13C over 22 years was about 5 per thousand, ranging from -27 to -22.3 per thousand. The most obvious pattern was a sharp decline in delta13C during latewood formation and a rapid increase with spring early growth. We conclude that the LA-C-GC-IRMS method will be useful in elucidating short-term climate effects on the delta13C signal in tree rings.

Carbon Isotopes↗

Regulation of nitrate uptake at the whole-tree level: interaction between nitrogen compounds, cytokinins and carbon metabolism.

Pedospheric nitrate uptake is closely integrated with the nitrogen (N) status and demand of the whole tree. Signaling substances communicating the N demand of the shoot to the roots are required in an integrated regulatory system. Besides phloem mobility, such signal compounds must have the potential to repress or increase nitrate uptake either at the transcriptional or post-transcriptional level. Amino compounds cycling within the tree are involved in the regulation of nitrate uptake. In many tree species, inorganic N is generally assimilated in roots, and amino acids--the direct products of N assimilation--are transported in the xylem to the sites of N demand. If the quantity of amino acids transported to the above-ground parts of the tree exceeds shoot N demand, some amino compounds are reallocated to the roots by phloem transport. Particular amino compounds exert transcriptional and post-transcriptional control over nitrate uptake by roots. Induction of nitrate transporters is mediated by nitrate or nitrite, or both, and possibly also by cytokinins, which cycle within the tree and act as both root-to-shoot and shoot-to-root signals. This review focuses on tree-specific requirements for N regulation and signaling, as well as the link between carbon metabolism and nitrate uptake.

Carbohydrate Metabolism↗

Variable responses of the depth of tree nitrogen uptake to pruning and competition.

Trees in cropped fields may improve nitrogen (N) use efficiency by intercepting leached N, but crop yield will be reduced if the trees compete strongly with crops for N. Ideal trees for intercropping will take up N from deeper soil layers not accessed by the crop species. Spatiotemporal aspects of tree nitrogen capture niches were investigated within a hedgerow intercropping system by placing 15N at three depths and monitoring 15N uptake by trees pruned either 25 or 4 days before application of 15N. Trees with contrasting rooting patterns (Gliricidia sepium L. and Peltophorum dasyrrachis (Miq.) Kurz) were grown in mixed hedgerows and intercropped with maize (Zea mays L.). Neither species showed significant N uptake during the 5-14 days after pruning, even though some shoot regrowth occurred during this time. Mean topsoil (0-5 cm depth) root length density of G. sepium was 520% greater than that of P. dasyrrachis, whereas total root length (0-100 cm depth) of G. sepium was only 450% greater. On average, G. sepium recovered 15 times as much 15N as P. dasyrrachis, following application of 15N at 5 cm depth, but the two species recovered a similar amount following application at 80 cm depth, suggesting that P. dasyrrachis had better niche complementarity with shallow rooting crops. However, both species showed strong plasticity in vertical N uptake pattern in response to competition from establishing maize plants. The species differed in their response: N uptake activity by G. sepium shifted down the soil profile in response to increasing competition from a growing maize crop (uptake from 80 cm depth changed from 4 to 9% of uptake from 5 cm depth), whereas N uptake by P. dasyrrachis became relatively shallow (uptake from 80 cm depth changed from 305 to 25% of uptake from 5 cm depth). Niche avoidance and increased competitiveness within the topsoil represent alternative responses to competition. The response displayed may be related to soil fertility in the species' natural habitats.

Adaptation, Physiological↗

Trees as indicators of subterranean water flow from a retired radioactive waste disposal site.

Tree sampling helped locate a subterranean flow of tritiated water from a low-level radioactive waste disposal site that had not been detected by well water monitoring alone. Deciduous trees growing in a natural forest on the hillsides downslope from the site were sampled for the presence of tritiated water in sap of maple trees and in leaf water extracted from oak and hickory trees. Elevated concentrations of 3H were detected in the leaf water extracted from several trees located 50 m downslope from the western boundary of the fenced exclusion zone. A 3-m-deep well drilled near these trees indicated that the source of tritiated water was a narrow zone of subterranean flow.

Radioactive Waste↗

Automated decision tree classification of corneal shape.

PURPOSE: The volume and complexity of data produced during videokeratography examinations present a challenge of interpretation. As a consequence, results are often analyzed qualitatively by subjective pattern recognition or reduced to comparisons of summary indices. We describe the application of decision tree induction, an automated machine learning classification method, to discriminate between normal and keratoconic corneal shapes in an objective and quantitative way. We then compared this method with other known classification methods. METHODS: The corneal surface was modeled with a seventh-order Zernike polynomial for 132 normal eyes of 92 subjects and 112 eyes of 71 subjects diagnosed with keratoconus. A decision tree classifier was induced using the C4.5 algorithm, and its classification performance was compared with the modified Rabinowitz-McDonnell index, Schwiegerling's Z3 index (Z3), Keratoconus Prediction Index (KPI), KISA%, and Cone Location and Magnitude Index using recommended classification thresholds for each method. We also evaluated the area under the receiver operator characteristic (ROC) curve for each classification method. RESULTS: Our decision tree classifier performed equal to or better than the other classifiers tested: accuracy was 92% and the area under the ROC curve was 0.97. Our decision tree classifier reduced the information needed to distinguish between normal and keratoconus eyes using four of 36 Zernike polynomial coefficients. The four surface features selected as classification attributes by the decision tree method were inferior elevation, greater sagittal depth, oblique toricity, and trefoil. CONCLUSION: Automated decision tree classification of corneal shape through Zernike polynomials is an accurate quantitative method of classification that is interpretable and can be generated from any instrument platform capable of raw elevation data output. This method of pattern classification is extendable to other classification problems.

Cornea↗