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Twilight-zone and canopy shade induction of the Athb-2 homeobox gene in green plants.

We present evidence that a novel phytochrome (other than phytochromes A and B, PHYA and PHYB) operative in green plants regulates the "twilight-inducible" expression of a plant homeobox gene (Athb-2). Light regulation of the Athb-2 gene is unique in that it is not induced by red (R)-rich daylight or by the light-dark transition but is instead induced by changes in the ratio of R to far-red (FR) light. These changes, which normally occur at dawn and dusk (end-of-day FR), also occur during the daytime under the canopy (shade avoidance). By using pure light sources and phyA/phyB null mutants, we demonstrated that the induction of Athb-2 by changes in the R/FR ratio is mediated for the most part by a novel phytochrome operative in green plants. Furthermore, PHYB plays a negative role in repressing the accumulation of Athb-2 mRNA in the dark and a minor role in the FR response. The strict correlation of Athb-2 expression with FR-induced growth phenomena suggests a role for the Athb-2 gene in mediating cell elongation. This interpretation is supported by the finding that the Athb-2 gene is expressed at high levels in rapidly elongating etiolated seedlings. Furthermore, as either R or FR light inhibits cell elongation in etiolated tissues, they also down-regulate the expression of Athb-2 mRNA. Thus, these data support the notion that changes in light quality perceived by a novel phytochrome regulate plant development through the action of the Athb-2 homeobox gene.

Journal Article↗

WIMOVAC: a software package for modelling the dynamics of plant leaf and canopy photosynthesis.

The ability to predict net carbon exchange and production of vegetation in response to predicted atmospheric and climate change is critical to assessing the potential impacts of these changes. Mathematical models provide an important tool in the study of whole plant, canopy and ecosystem responses to global environmental change. Because this requires prediction beyond experience, mechanistic rather than empirical models are needed. The uniformity and strong understanding of the photosynthetic process, which is the primary point of response of plant production to global atmospheric change, provides a basis for such an approach. Existing modelling systems have been developed primarily for expert modellers and have not been easily accessible to experimentalists, managers and students. Here we describe a modular modelling system operating within Windows to provide this access. WIMOVAC (Windows Intuitive Model of Vegetation response to Atmosphere and Climate Change) is designed to facilitate the modelling of various aspects of plant photosynthesis with particular emphasis on the effects of global climate change. WIMOVAC has been designed to run on IBM PC-compatible computers running Microsoft Windows. The package allows the sophisticated control of the simulation processes for photosynthesis through a standardized Windows user interface and provides automatically formatted results as either tabulated data or as a range of customizable graphs. WIMOVAC has been written in Microsoft Visual Basic, to facilitate the rapid development of user-friendly modules within the familiar Windows framework, while allowing a structured development. The highly interactive nature of controls adopted by WIMOVAC makes it suitable for research, management and educational purposes.

Algorithms↗

Similar reproductive status and body size of horse flies (Diptera: Tabanidae) attracted to carbon dioxide-baited canopy traps and a Jersey bullock.

The reproductive status and body size of four Tabanus species collected from canopy traps baited with carbon dioxide and from a Jersey bullock were compared. Parity rates, sperm prevalence, stage of follicular development in terminal follicles of parous females, prevalence of females retaining eggs, average number of eggs retained in parous flies, and the body size of parous females did not differ significantly between sampling methods. Based on the presence of nulliparous host-seeking flies, Tabanus pallidescens Philip and T. wilsoni Pechuman can be added to the list of tabanids found to be anautogenous.

Animals↗

Cytogenetic and isozyme profile of Sabethes cyaneus: a mosquito of the neotropical canopy.

Sabethes cyaneus, a newly colonized culicine mosquito from the Panamanian forest canopy, has a distribution range from Honduras to Argentina. Cytogenetic studies, the first on any Sabethes species, revealed a karyotype of three pairs of homomorphic chromosomes (2n = 6). Well-developed polytene chromosomes were discovered in the larval salivary glands and a photomap standard was constructed. Clear banding patterns and consistent landmarks distinguished each of the six arms. Substantial asynapsis occurred in the three polytene chromosomes, although the banding pattern of the homologous regions appeared homosequential. A nucleolar organizer was localized in region 5A of the shortest chromosome by the recurrent association of this region with the nucleolus. The large band in region 5A was found heterozygous for width and a deletion. Additional, less conspicuous, polymorphisms for band width and staining intensity were distributed throughout the genome. Biochemical studies of 31 enzyme loci revealed 10 loci to be polymorphic, with an average heterozygosity of 13 percent. Differential expression in developmental stages occurred for 11 loci involving six enzymes.

Animals↗

Root growth and carbohydrate responses in bearing citrus trees following partial canopy removal.

In August, eight 4-m tall citrus trees were pruned by removing the top third of their canopy. Eight unpruned trees served as controls. Root growth, which was examined nondestructively with minirhizotrons over a four-month period, tended to be less in the pruned than unpruned trees seven days after pruning and this difference was significant (P < 0.05) from 14 to 49 days after pruning. Total reducing and ketone sugars (includes free fructose, sucrose and fructans) in the fine roots were less in pruned than unpruned trees 20 days after pruning, but not thereafter. By 30 days after pruning, at least 20% of the roots of the pruned trees at a soil depth of 9 to 35 cm apparently died. By 63 days after pruning, root length density had recovered to that of the unpruned trees, although starch reserves were 18% less in the fine roots of pruned than unpruned trees at this time. Nine to eleven months after pruning (May to July), total biomass of leaves and fine roots to a depth of 1 m were similar in pruned and unpruned trees. However, fruit biomass harvested in April from pruned trees was only 24% of that in the unpruned trees. In May, nonstructural carbohydrates in the fine and coarse roots of pruned trees were generally greater than in unpruned trees, possibly reflecting previous differences in fruit production.

Journal Article↗

Patterns of hydraulic architecture and water relations of two tropical canopy trees with contrasting leaf phenologies: Ochroma pyramidale and Pseudobombax septenatum.

Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.

Journal Article↗

Interactive effects of atmospheric carbon dioxide and leaf nitrogen concentration on canopy light use efficiency: a modeling analysis.

Potential increases in plant productivity in response to increasing atmospheric CO(2) concentration are likely to be constrained by nutrient limitations. However, the interactive effects of nitrogen nutrition and CO(2) concentration on growth are difficult to define because both factors affect several aspects of growth, including photosynthesis, respiration, and leaf area. By expressing growth as a product of light intercepted and light use efficiency (epsilon), it is possible to decouple the effects of nutrient availability and CO(2) concentration on photosynthetic rates from their effects on other aspects of plant growth. I used measured responses of leaf photosynthesis to leaf nitrogen (N) content and CO(2) concentration to parameterize a model of canopy radiation absorption and photosynthesis, and then used the model to estimate the response of epsilon to elevated CO(2) concentration for Pinus radiata D. Don, Nothofagus fusca (Hook. f.) Ørst. and Eucalyptus grandis W. Hill ex Maiden. Down-regulation of photosynthesis at elevated CO(2) was represented as a reduction in either leaf N content or leaf Rubisco activity. The response of epsilon to elevated CO(2), which differed among the three species, was analyzed in terms of the underlying relationships between leaf photosynthesis and leaf N content. The response was independent of leaf N content when photosynthesis was down-regulated to the same extent at low and high leaf N content. Interactive effects of N availability and CO(2) on growth are thus likely to be the result of either differences in down-regulation of photosynthesis at low and high N availability or interactive effects of CO(2) and N availability on other aspects of plant growth.

Journal Article↗

Response of giant sequoia canopy foliage to elevated concentrations of atmospheric ozone.

We examined the physiological response of foliage in the upper third of the canopy of 125-year-old giant sequoia (Sequoiadendron giganteum Buchholz.) trees to a 61-day exposure to 0.25x, 1x, 2x or 3x ambient ozone concentration. Four branch exposure chambers, one per ozone treatment, were installed on 1-m long secondary branches of each tree at a height of 34 m. No visible symptoms of foliar ozone damage were apparent throughout the 61-day exposure period and none of the ozone treatments affected branch growth. Despite the similarity in ozone concentrations in the branch chambers within a treatment, the trees exhibited different physiological responses to increasing ozone uptake. Differences in diurnal and seasonal patterns of g(s) among the trees led to a 2-fold greater ozone uptake in tree No. 2 compared with trees Nos. 1 and 3. Tree No. 3 had significantly higher CER and g(s) at 0.25x ambient ozone than trees Nos. 1 and 2, and g(s) and CER of tree No. 3 declined with increasing ozone uptake. The y-intercept of the regression for dark respiration versus ozone uptake was significantly lower for tree No. 2 than for trees Nos. 1 and 3. In the 0.25x and 1x ozone treatments, the chlorophyll concentration of current-year foliage of trees Nos. 1 and 2 was significantly higher than that of current-year foliage of tree No. 3. Chlorophyll concentration of current-year foliage on tree No. 1 did not decline with increasing ozone uptake. In all trees, total needle water potential decreased with increasing ozone uptake, but turgor was constant. Although tree No. 2 had the greatest ozone uptake, g(s) was highest and foliar chlorophyll concentration was lowest in tree No. 3 in the 0.25x and 1x ambient atmospheric ozone treatments.

Journal Article↗

Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan.

Stem length and leaf area of current-year shoots were measured in saplings of eight broad-leaved evergreen tree species growing under a forest canopy. Stem length varied over a range of one to two orders of magnitude within each species. In all species, both the number of leaves and the mean stem length between successive leaves were greater in longer shoots. Mean leaf size and stem length were not correlated in six of eight species, and only weakly positively correlated in the other two species. Thus, total leaf area per stem increased with stem length, but not in direct proportion: leaf area per stem length was smaller in shoots with long stems and larger in shoots with short stems. I conclude that the within-species variation in the leaf-stem balance of current-year shoots is related to variation in shoot functional roles, as has been observed for long and short shoots in many deciduous tree species: shoots with long stems are extension oriented and contribute to the framework of the crown, whereas shoots with short stems serve mainly for leaf display. Among species, large differences were found in the leaf area per stem length ratio. In the species with larger leaf area per stem length ratios, leaves had narrower blades or longer petioles, or both, resulting in a reduction of mutual shading among the leaves on the shoot.

Journal Article↗

Limitation of leaf carbon gain by stomatal and photochemical processes in the top canopy of Macaranga conifera, a tropical pioneer tree.

Diurnal changes in gas exchange and chlorophyll fluorescence were measured in the top canopy leaves of the tropical rainforest tree species, Macaranga conifera (Zoll.) Muell. Arg. during a drought year. Maximum values of net photosynthetic rate (P(n), 10 &mgr;mol m(-2) s(-1)) and stomatal conductance (g(s), 0.2 mol m(-2) s(-1)) were found in east-facing leaves in early morning. After 1000 h, both P(n) and g(s) decreased. Minimum daytime values of P(n), g(s), and photosystem II (PSII) quantum yield (DeltaF/F(m)') were found in horizontally fixed leaves. At a given electron transport rate through PSII (ETR), P(n) was higher in early morning than at midday, suggesting a high rate of photorespiration at midday. We tested the hypothesis that the effect of low leaf temperature (T(leaf)) on P(n) is significant in the early morning, whereas the effect of low g(s) on P(n) predominates at midday. In the early morning, when T(leaf) was increased from 32 to 38 degrees C by artificial heating, P(n) at a given ETR decreased 29%, suggesting that the low T(leaf) was associated with a high P(n). When T(leaf) at midday was decreased from 37 to 32 degrees C by artificial cooling, P(n) increased 22%, but P(n) at a given ETR was higher in early morning than at midday, even at the same low T(leaf) (32 degrees C). This suggests that the rate of photorespiration was higher at midday than in early morning because low g(s) at midday caused a reduction in leaf intercellular CO(2) concentration. We conclude that low P(n) at midday was the result of both a reduction in the photochemical process and an increase in stomatal limitation.

Journal Article↗

Regulation of water flux through tropical forest canopy trees: do universal rules apply?

Tropical moist forests are notable for their richness in tree species. The presence of such a diverse tree flora presents potential problems for scaling up estimates of water use from individual trees to entire stands and for drawing generalizations about physiological regulation of water use in tropical trees. We measured sapwood area or sap flow, or both, in 27 co-occurring canopy species in a Panamanian forest to determine the extent to which relationships between tree size, sapwood area and sap flow were species-specific, or whether they were constrained by universal functional relationships between tree size, conducting xylem area, and water use. For the 24 species in which active xylem area was estimated over a range of size classes, diameter at breast height (DBH) accounted for 98% of the variation in sapwood area and 67% of the variation in sapwood depth when data for all species were combined. The DBH alone also accounted for > or = 90% of the variation in both maximum and total daily sap flux density in the outermost 2 cm of sapwood for all species taken together. Maximum sap flux density measured near the base of the tree occurred at about 1,400 h in the largest trees and 1,130 h in the smallest trees studied, and DBH accounted for 93% of the variation in the time of day at which maximum sap flow occurred. The shared relationship between tree size and time of maximum sap flow at the base of the tree suggests that a common relationship between diurnal stem water storage capacity and tree size existed. These results are consistent with a recent hypothesis that allometric scaling of plant vascular systems, and therefore water use, is universal.

Panama↗

Effects of thinning on wood production, leaf area index, transpiration and canopy interception of a plantation subject to drought.

We conducted thinning trials in a 5-year-old Eucalyptus globulus ssp. globulus Labill plantation near Warrenbayne, northeastern Victoria, Australia, where soil salinization and waterlogging are common, and assessed treatment effects on tree growth, water use and survival. Half-hectare plots were thinned from the original density of 1100 stems ha(-1) to densities of 800, 600 and 400 stems ha(-1), and stem diameter increment, leaf area index, transpiration, canopy interception and depth of tree water source monitored for 21 months. Two drought periods occurred during the study, rainfall was 30% below the long-term average and there was severe mortality in all three plots. Analysis of deuterium abundance in soil and xylem water indicated that the trees accessed water only from the top meter of the soil profile. Transpiration rates were higher in the most heavily thinned plot than in the least thinned plot, which underwent a reduction in basal area during the study. The most heavily thinned plot increased in basal area by 10% during the study. Edge trees had significantly greater diameters than trees from the middle of the plots.

Disasters↗

Mean canopy stomatal conductance responses to water and nutrient availabilities in Picea abies and Pinus taeda.

We compared sap-flux-scaled, mean, canopy stomatal conductance (GS) between Picea abies (L.) Karst. in Sweden and Pinus taeda (L.) in North Carolina, both growing on nutritionally poor soils. Stomatal conductance of Picea abies was approximately half that of Pinus taeda and the sensitivity of GS in Picea abies to vapor pressure deficit (D) was lower than in Pinus taeda. Optimal fertilization increased leaf area index (L) two- and threefold in Pinus taeda and Picea abies, respectively, regardless of whether irrigation was increased. Although it increased L, fertilization did not increase GS in Picea abies unless irrigation was also provided. In Pinus taeda growing on coarse, sandy soils, the doubling of L in response to fertilization reduced GS sharply unless irrigation was also provided. The reduction in GS with fertilization in the absence of irrigation resulted from the production of fine roots with low saturated hydraulic conductivity. When Pinus taeda received both fertilization and irrigation, the increase in L was accompanied by a large increase in GS. In Pinus taeda, a reference GS (defined as GS at D = 1 kPa; GSR) decreased in all treatments with decreasing volumetric soil water content (theta). In Picea abies, theta varied little within a treatment, but overall, GSR declined with theta, reaching lowest values when drought was imposed by the interception of precipitation. Despite the large difference in GS both between Picea abies and Pinus taeda and among treatments, stem growth was related to absorbed radiation, and stem growth response to treatment reflected mostly the changes in L.

Biomass↗

Canopy structure and light interception in Quercus petraea seedlings in relation to light regime and plant density.

Foliage structure was measured on 1- and 2-year-old Quercus petraea (Matt.) Liebl. seedlings grown in 100 or 18% sunlight at a planting density of 2.8 or 25 plants per m(2). A three-dimensional digitizing device was used to acquire the spatial position and orientation of all leaves within the seedlings and of all seedlings within the plot. The data were used to obtain (1) quantitative information on canopy structure, including leaf area index (LAI), seedling leaf area, number of leaves, leaf area density and leaf orientation; and (2) structural information on foliage arrangement from virtual images to estimate light interception by individual seedlings (STAR) and light partitioning among seedlings. During the second year, shading significantly reduced total leaf area and number of leaves but increased individual leaf area. The STAR was greater for seedlings in shade than in full sunlight because of the more horizontal orientation of leaves. Leaf area density was unaffected by the full sun treatments, and changes in leaf area dispersion had no effect on light-interception efficiency. No plant density effect was observed during the first year. During the second year, only the high plant density treatment induced mutual shading between seedlings, resulting in greater competition for light among seedlings in the full sun treatment than in the shade treatment. The small treatment-induced changes in light interception indicate that Q. petraea has low morphological plasticity of foliage structure compared with other species.

Light↗

Dependence of needle architecture and chemical composition on canopy light availability in three North American Pinus species with contrasting needle length.

Morphology and chemical composition of needles of shade-intolerant southern conifers (Pinus palustris Mill. (mean needle length +/- SD = 29.1 +/- 4.1 cm), P. taeda L. (12.3 +/- 2.9 cm) and P. virginiana Mill. (5.1 +/- 0.8 cm)) were studied to test the hypothesis that foliage acclimation potential to canopy light gradients is generally low for shade-intolerant species, and in particular, because of mechanical limitations, in species with longer needles. Plasticity for each needle variable was defined as the slope of the foliar characteristic versus irradiance relationship. A novel geometrical model for needle area and volume calculation was employed for the three-needled species P. palustris and P. taeda. Needle thickness (T) strongly increased, but width (W) was less variable with increasing daily integrated quantum flux density averaged over the season (Q(int)), resulting in changes in cross-sectional needle shape that were manifested in a positive relationship between the total to projected needle area ratio (A(T)/A(P)) and Q(int) in the three-needled species. In contrast, cross-sectional needle geometry was only slightly modified by irradiance in the two-needled conifer P. virginiana. Needle dry mass per unit total needle area (M(T)) was positively related to Q(int) in all species, leading to greater foliar nitrogen contents per unit area at higher irradiances. Separate examination of the components of M(T) (density (D) and the volume (V) to A(T) ratio; M(T) = DV/A(T)) indicated that the positive effect of light on M(T) resulted solely from increases in V/A(T), i.e., from increases in the thickness of foliage elements. Foliar chlorophyll content per unit mass increased with increasing Q(int), allowing an improvement in light-harvesting efficiency in low light. The variables characterizing needle material properties (D, the dry to fresh mass ratio, and needle carbon content per unit mass) were generally independent of Q(int), suggesting that needles were less stiff and had greater tip deflections under their own weight at lower irradiances because of smaller W and T. Comparisons with the literature revealed that plasticity in foliar characteristics tended to be lower in the studied shade- intolerant species than in shade-tolerant conifers, but plasticity among the investigated species was unaffected by needle length. However, we argue that, because of mechanical limitations, plastic changes in needle cross section in response to low irradiance may decrease rather than increase light-interception efficiency in long-needled species.

Biomass↗

Changes in shoot allometry with increasing tree height in a tropical canopy species, Elateriospermum tapos.

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.

Biomass↗

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees.

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.

Chlorophyll↗