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Why is it advantageous for animals to detect celestial polarization in the ultraviolet? Skylight polarization under clouds and canopies is strongest in the UV.

The perception of skylight polarization in the ultraviolet (UV) by many insect species for orientation purposes is rather surprising, because both the degree of linear polarization and the radiance of light from the clear sky are considerably lower in the UV than in the blue or green. In this work we call this the "UV-sky-pol paradox". Although in the past, several attempts have been made to resolve this paradox, none of them was convincing. We present here a possible quantitative resolution to the paradox. We show by a model calculation that if the air layer between a cloud and a ground-based observer is partly sunlit, the degree of linear polarization p of skylight originating from the cloudy region is highest in the UV, because in this spectral range the unpolarized UV-deficient cloudlight dilutes least the polarized light scattered in the air beneath the cloud. Similarly, if the air under foliage is partly sunlit, p of downwelling light from the canopied region is maximal in the UV, because in this part of spectrum the unpolarized UV-deficient green canopylight dilutes least the polarized light scattered in the air beneath the canopy. Therefore, the detection of polarization of downwelling light under clouds or canopies is most advantageous in the UV, in which spectral range the risk is the smallest that the degree of polarization p is lower than the threshold p(tr) of polarization sensitivity in animals. On the other hand, under clear skies there is no favoured wavelength for perception of celestial polarization, because p of skylight is high enough (p > p(tr)) at all wavelengths. We show that there is an analogy between the detection of UV skylight polarization and the polarotactic water detection in the UV. However, insects perceive skylight polarization by UV or blue or green receptors. The question, why they differ in the spectral channel used for the detection of celestial polarization cannot be answered at the present time, because data are insufficient. Nevertheless, we present here one possible atmospheric optical reason why certain visual systems involved in detecting celestial polarization, are specifically tuned to the UV part of the spectrum.

Animals↗

Plant population growth and competition in a light gradient: a mathematical model of canopy partitioning.

Can a difference in the heights at which plants place their leaves, a pattern we call canopy partitioning, make it possible for two competing plant species to coexist? To find out, we examine a model of clonal plants living in a nonseasonal environment that relates the dynamical behavior and competitive abilities of plant populations to the structural and functional features of the plants that form them. This examination emphasizes whole plant performance in the vertical light gradient caused by self-shading. This first of three related papers formulates a prototype single species Canopy Structure Model from biological first principles and shows how all plant properties work together to determine population persistence and equilibrium abundance. Population persistence is favored, and equilibrium abundance is increased, by high irradiance, high maximum photosynthesis rate, rapid saturation of the photosynthetic response to increased irradiance, low tissue respiration rate, small amounts of stem and root tissue necessary to support the needs of leaves, and low density of leaf, stem, and root tissues. In particular, equilibrium abundance decreases as mean leaf height increases because of the increased cost of manufacturing and maintaining stem tissue. All conclusions arise from this formulation by straightforward analysis. The argument concludes by stating this formulation's straightforward extension, called a Canopy Partitioning Model, to two competing species.

Ecosystem↗

Effects of herbivorous snails and macroalgal canopy on recruitment and early survivorship of the barnacle Semibalanus balanoides (L.).

This study investigated the effects of grazing by the herbivorous snail Littorina littorea (L.) and the presence of a macroalgal canopy on recruitment and early survivorship of the barnacle Semibalanus balanoides (L.) at four sites of various distances from the mouth of Toothacher Cove on Swans Island in the Gulf of Maine in 1998. Recruitment plates were attached to the substrate at each site in all possible treatment combinations, including either presence of absence of macroalgal canopy, presence or absence of the herbivore, and presence or absence of a cage. Significant differences in recruitment were observed among sites in April and May, but not in June. The average number of recruits also varied significantly among herbivore treatments in May and June, and in the May macroalgaexherbivore interaction. Survivorship did not differ among sites, but did vary significantly among herbivore treatments. These differences were due almost entirely to consistently low recruitment and survival in the uncaged or open controls. Comparisons of the caged and open treatments explained between 88 and 99% of the variation in the respective main effects tests. The presence or absence of L. littorea or of macroalgal canopy had little effect on S. balanoides recruitment or survivorship. Results support previous conclusions that recruitment is greater on more exposed shores and suggest that exclusion cages may protect barnacle recruits from factors such as desiccation and the whiplash effect of macroalgal fronds.

Journal Article↗

The effects of seagrass (Zostera japonica) canopy structure on associated fauna: a study using artificial seagrass units and sampling of natural beds.

The importance of seagrass canopy to associated fauna was assessed by comparing the species richness, abundance and diversity of the epi- and infaunal macroinvertebrate assemblages in a seagrass (Zostera japonica Ascherson and Graebner) bed and the adjacent unvegetated area in Hong Kong. Seagrass cover had significant effects on the composition and abundance of the associated fauna and the amount of detritus accumulated on the sediment surface. Detritus abundance was significantly higher in the seagrass bed, and was positively correlated with both the above- and belowground biomass of Z. japonica. Both the abundance and species richness of the epi- and infauna were significantly positively correlated with the belowground biomass of the seagrass and detritus standing crop. Macrofaunal species richness was higher (118) in the seagrass bed than the adjacent unvegetated areas (70), with a higher degree of similarity between the infauna than the epifauna of the two habitats. While all species recorded from the unvegetated areas were found in the seagrass bed, 48 species occurred only in the seagrass-covered areas. Species richness of epifauna was significantly higher in the seagrass bed, but there was no difference between infaunal species of the two habitats. On the contrary, faunal (epi- and infauna) abundance was significantly higher in seagrass areas. The seagrass bed also supported species of small tellinid bivalves previously not recorded from Hong Kong. Artificial seagrass units (ASUs, 0.2 m(2)) with four combinations of leaf density and leaf length and a control (bare sand) were placed at short distances from natural patches of Z. japonica. The composition, abundance and biomass of the epibenthos associated with the ASUs and the control were recorded after 3 months in the field. While species richness did not differ among the treatments, total abundance of epibenthos was significantly higher in the high density-long leaves (HL) treatment than in the control. Results of a discriminant analysis using log-transformed abundance data suggest that the gastropod Clithon oualaniensis, the mussel Musculista senhousia and the crab Thalamita sp. were important species distinguishing the assemblages in the various treatments. All the three species were significantly more abundant in the HL treatment than in the low density-short leaves (LS) treatment and the control. By contrast, there was no significant difference in the biomass of the epifauna, but discriminant analysis again separated the five treatments based on the composition of the biomass, with the same three species identified as the most important discriminating species. The species richness and abundance of the epifauna associated with the ASUs were similar to the adjacent unvegetated areas, but significantly lower than in the Zostera patches. The physical canopy structure of Z. japonica beds increased the abundance of the epibenthos, potentially through provision of canopy and indirectly through trapping of detritus.

Journal Article↗

Modeling above-canopy CO2 flux and evapotranspiration in wheat.

Simulations of above-canopy water vapor and CO2 fluxes were calculated by the USGF linked model of canopy gas exchange and subsurface processes for the 1996-1997 winter wheat season at the AmeriFlux Wheat Site, Oklahoma. Soil surface CO2 flux plus canopy gas exchange and transpiration plus soil evaporation modeled the CO2 and water vapor fluxes, respectively. Parameter values for net photosynthesis, respiration and transpiration were obtained from published sources, generated from Wheat Site data, or estimated by minimizing standard deviation between model and data. The mean measured downward flux of CO2 during rapid growth and maturity of the crop was -0.45 mg m(-2) s(-1) compared to simulated flux of -0.47. Simulated downward CO2 flux exceeded measured values during rapid growth of the crop but underestimated the flux during maturity. For the entire 285-day period, the mean measured upward CO2 flux at night was 0.06 and simulated flux was 0.05.

Carbon Dioxide↗

Importance of the forest canopy to fluxes of methyl mercury and total mercury to boreal ecosystems.

The forest canopy was an important contributor to fluxes of methyl mercury (MeHg) and total mercury (THg) to the forest floor of boreal uplands and wetlands and potentially to downstream lakes, at the Experimental Lakes Area (ELA), northwestern Ontario. The estimated fluxes of MeHg and THg in throughfall plus litterfall below the forest canopy were 2 and 3 times greater than annual fluxes by direct wet deposition of MeHg (0.9 mg of MeHg ha(-1)) and THg (71 mg of THg ha(-1)). Almost all of the increased flux of MeHg and THg under the forest canopy occurred as litterfall (0.14-1.3 mg of MeHg ha(-1) yr(-1) and 110-220 mg of THg ha(-1) yr(-1)). Throughfall added no MeHg and approximately 9 mg of THg ha(-1) yr(-1) to wet deposition at ELA, unlike in other regions of the world where atmospheric deposition was more heavily contaminated. These data suggest that dry deposition of Hg on foliage as an aerosol or reactive gaseous Hg (RGM) species is low at ELA, a finding supported by preliminary measurements of RGM there. Annual total deposition from throughfall and litterfall under a fire-regenerated 19-yr-old jack pine/birch forest was 1.7 mg of MeHg ha(-1) and 200 mg of THg ha(-1). We found that average annual accumulation of MeHg and THg in the surficial litter/fungal layer of soils since the last forest fire varied between 0.6 and 1.6 mg of MeHg ha(-1) and between 130 and 590 mg of THg ha(-1) among sites differing in drainage and soil moisture. When soil Hg accumulation sites were matched with similar sites where litterfall and throughfall were collected, measured fluxes of THg to the forest floor (sources) were similar to our estimates of longterm soil accumulation rates (sinks), suggesting that the Hg in litterfall and throughfall is a new and not a recycled input of Hg to forested ecosystems. However, further research is required to determine the proportion of Hg in litterfall that is being biogeochemically recycled within forest and wetland ecosystems and, thus, does not represent new inputs to the forest ecosystem.

Aerosols↗

Accumulation of persistent organic pollutants in canopies of different forest types: role of species composition and altitudinal-temperature gradient.

Leaves from the dominant tree species in three different alpine forests were sampled along an altitudinal gradient and analyzed for HCB, alpha- and gamma-HCH, and PCBs. The mean canopy concentration was calculated, considering the relative abundance of each species in the respective forest. Compound fractionation occurred in the vegetation along the altitudinal/temperature gradient. Results were compared with air concentrations and in-field plant/air partition coefficients (K(PA)) were calculated for each species; this showed differences between broadleaves and needles. The mean canopy/air partition coefficient (K(CA)) was also calculated by averaging results from single species. The variability of persistent organic pollutants distribution in canopies is discussed considering two main factors, the altitudinal/temperature gradient and the species composition. The latter is responsible for most of the concentration variability of the more volatile compounds. A model to calculate dry gaseous deposition to different forest canopies is presented.

Air Pollutants↗

Doubling the estimate of invertebrate biomass in a rainforest canopy.

Forest canopies represent the functional interface between 90% of the Earth's terrestrial biomass and the atmosphere and include some of the most threatened of all terrestrial ecosystems. However, we lack even a basic understanding of how the biomass of plants and animals is distributed throughout forest canopies, even though this information is vital for estimating energy flow, carbon cycling, resource use and the transfer of materials within this ecosystem. Here we measure the biomass of invertebrates living in a common rainforest epiphyte, describe a striking relationship between fern size and the biomass of animals within the ferns, and reveal that one large epiphyte may contain an invertebrate biomass similar to that found in the whole of the rest of the tree crown on which it is growing. Using these data, we show that including the fauna of these epiphytes--a neglected component in rainforest ecosystems--can more than double our estimate of the total invertebrate biomass in an entire rainforest canopy.

Animals↗

Birds defend trees from herbivores in a Neotropical forest canopy.

Most forest birds include arthropods in their diet, sometimes specializing on arthropods that consume plant foliage. Experimental tests of whether bird predation on arthropods can reduce plant damage, however, are few and restricted to relatively low-diversity systems. Here, we describe an experimental test in a diverse tropical forest of whether birds indirectly defend foliage from arthropod herbivores. We also compare how the indirect effects of bird predation vary with different levels of foliage productivity in the canopy vs. the understory. For three Neotropical tree species, we observed that birds decreased local arthropod densities on canopy branches and reduced consequent damage to leaves. In contrast, we observed no evidence of bird-arthropod limitation on conspecific saplings in the less productive understory of the same forest. Our results support theory that predicts trophic cascades where productivity is high and suggest that birds play an important role in Neotropical communities by means of their indirect defense of some canopy tree species.

Journal Article↗

The alpha-tocopherol content of leaves of pedunculate oak (Quercus robur L.)--variation over the growing season and along the vertical light gradient in the canopy.

This study was performed in order to investigate whether the actual requirement for defence against photo-oxidative stress is reflected by the alpha-tocopherol (alpha-Toco) content in leaves of pedunculate oak (Quercus robur L.). Antioxidants and pigments were quantified in leaves that were collected on six days between May and September 2000 in a mixed pine/oak forest at canopy positions differing in light environment. Pools of hydrophilic antioxidants and photo-protective xanthophyll cycle pigments (V + A + Z) reflected the anti-oxidative demand, as these pools increased with the average light intensity to which the leaves were acclimated. The photo-protective demand was not the determinant of the alpha-Toco content of oak leaves, as (1) foliage of a young oak, exposed to low light levels in the understorey, contained higher amounts of this lipophilic antioxidant than leaves sampled from semimature oaks at canopy positions with a similar light environment, and (2) a strong increase in the alpha-Toco content over the growing season was detected at each investigated crown position, whereas the V + A + Z pool did not show a concomitant accumulation during leaf ageing. The rate of alpha-Toco accumulation differed distinctly between samples taken at different canopy positions.

Acclimatization↗

Leaf and canopy photosynthetic characteristics of cotton (Gossypium hirsutum) under elevated CO2 concentration and UV-B radiation.

Increases in both atmospheric CO2 concentration ([CO2]) and ultraviolet-B (UV-B) radiation on the Earth's surface are features of current climate change patterns. An experiment was conducted in sunlit, controlled environment chambers known as Soil-Plant-Atmosphere-Research (SPAR) units to determine interactive effects of elevated [CO2] and UV-B radiation on leaf and canopy photosynthetic characteristics of cotton. Six treatments were comprised of two CO2 levels of 360 (ambient) and 720 (elevated) microL L(-1) and three levels of 0 (control), 8, and 16 kJ m(-2) d(-1) biologically effective UV-B radiation. Treatments were imposed for 66 days from crop emergence through three weeks after the first flower stage. Plants grown in elevated [CO2] had significantly greater leaf area, higher leaf and canopy net photosynthetic rates (PN), lower dark respiration rate (Rd), and lower light compensation point (LCP) than plants grown in ambient [CO2]. There was no difference in CO2 compensation point (gamma), maximum rate of Rubisco activity (Vcmax), or light-saturated rate of electron transport (Jmax) between ambient and elevated CO2 treatments. When plants were grown in 8 kJ m(-2) d(-1) UV-B radiation, most of the measured photosynthetic parameters did not differ from control plants. High UV-B (16 kJ) radiation, however, caused 47-50% smaller leaf area, 38-44% lower leaf PN, 72-74% lower Vcmax, and 61-66% lower Jmax compared to the control. There were no interactive effects of [CO2] and UV-B radiation on most of the photosynthetic parameters measured. From the results, it is concluded that decreased canopy photosynthesis due to enhanced UV-B radiation in cotton is associated with both smaller leaf area and lower leaf PN, and loss of Rubisco activity and electron transport are two major factors in UV-B inhibition of leaf PN.

Carbon Dioxide↗

Canopy gradients in delta(18)O of organic matter as ecophysiological tool.

We determined vertical oxygen isotope gradients of leaf organic matter for a grassland in Switzerland and a mountain beech forest (Fagus sylvatica) in Northern Italy. A distinctly positive (18)O/(16)O gradient with height above ground was found for the grassland (7.9/1000 m(-1), p < 0.001), whereas the gradient was negative for the forest (-0.077/1000 m(-1), p < 0.001). The results are consistent with microclimatic measurements, although large isotope variations between the species have to be taken into account for the grassland. A conceptual scheme is shown which relates the isotope enrichment to the canopy density, considering the effects of transpiration and canopy structure. We conclude that the analysis of the within canopy variation in delta(18)O of organic matter can be used to provide long-term estimates of leaf water isotope composition, thus improving existing isotope methods to determine the gas-exchange between vegetation and atmosphere.

Atmosphere↗

Variability in Leaf Morphology and Chemical Composition as a Function of Canopy Light Environment in Coexisting Deciduous Trees.

Morphology, chemical composition, and photosynthetic capacity of leaf laminas were investigated in Populus tremula L. and Tilia cordata Mill. along a canopy light gradient. Variables determining the thickness of boundary layer for heat and water exchange at a given wind speed-effective leaf width (Ww) and length (Wd)-scaled positively with daily integrated quantum flux density averaged over the season (Qint, mol m-2 d-1) in T. cordata, but Wd decreased and Ww was constant with increasing Qint in P. tremula, bringing about a moderately improved capacity for convective cooling at greater irradiances in the latter species. Foliar stable carbon isotope discrimination (Delta) decreased with increasing Qint, demonstrating that, possibly because of more severe foliar water stress, leaves operated at a lower intercellular CO2 concentration in the upper canopy. Further analysis of foliar characteristics provided additional evidence of the interaction between water stress and Qint. Leaf dry matter content and both components of lamina dry mass per area (MA)-lamina thickness and density (dry mass per unit volume, rhoB)-increased with increasing Qint in both species. The rhoB and lamina dry matter content were also positively related to lamina carbon concentration, variability in which along the canopy was related to changes in carbon-rich lignin concentration. Since both increases in lamina density and lignin concentration improve leaf tolerance of low-water potentials, these foliar modifications were interpreted as indicative of acclimation to enhanced water limitations in high light. For the whole material, foliar nitrogen concentrations decreased with increasing rhoB, suggesting that an improvement of foliar mechanical strength may result in declining foliar assimilative potential. However, foliar photosynthetic electron transport capacity per unit area increased with increasing rhoB, possibly because increases in rhoB with light are not only attributable to greater cell wall lignification but also to denser packing of leaf cells, in particular, in fractional increases in palisade tissues with Qint. Because of a positive scaling of leaf thickness and density with total tree height, MA was greater in taller trees of T. cordata, foliage of which also had lower Delta and was likely to function with less open stomata. In summary, we conclude that leaf water stress, which scales with both Qint and total tree height, is a major factor altering foliage structure and assimilative capacity.

Journal Article↗

Acclimation of photosynthesis to light and canopy nitrogen distribution: an interpretation.

BACKGROUND AND AIMS: Acclimation of photosynthesis to light and its connection with canopy nitrogen (N) distribution are considered. An interpretation of a proportionality between light-saturated photosynthesis and local averaged leaf irradiance is proposed by means of a simple model. MODEL: The model assumes (a) local irradiance drives synthesis of photosynthetic protein from metabolic N; (b) photosynthetic N is slowly degraded over approx. 5-7 d; (c) metabolic N is equally available through the canopy. CONCLUSIONS: The kinetics of acclimation at different light levels may provide a way of parameterizing and testing the model. The model provides a rationale for the proportionality assumption mentioned above, which, while it is consistent with much experimental work, is valuable because it allows canopy photosynthesis to be calculated analytically.

Acclimatization↗

Species-specific variation in the importance of the spectral quality gradient in canopies as a signal for photosynthetic resource partitioning.

BACKGROUND AND AIMS: Plants adjust the distribution of photosynthetic capacity and chlorophyll to canopy density. The importance of the gradient in the red : far-red ratio (R : FR) relative to the irradiance gradient was studied for its perception with respect to this partitioning of photosynthetic resources. Whether the relative importance of these two signals varied between six species of different growth habit (Phaseolus vulgaris, Lysimachia vulgaris, Hedera helix, Ficus benjamina, Carex acutiformis and Brachypodium pinnatum) was investigated further. METHODS: Single leaves of plants were shaded in daylight by a spectrally neutral filter or a leaf. In another experiment, leaves were treated with supplemental FR. In most cases, treatment effects were evaluated after 2 weeks. KEY RESULTS: Nitrogen and photosynthetic capacity (Amax) per leaf area, parameters pertaining to between-leaf resource partitioning, were strongly reduced in neutral shade but not additionally by spectral leaf shade. Supplemental FR reduced these parameters also, except in Carex. Acceleration of induction of senescence was observed in spectral leaf shade in primary bean leaves. Amax per unit chlorophyll, a parameter pertaining to within-leaf resource partitioning, was reduced in neutral shade, but not in spectral leaf shade or supplemental FR. CONCLUSIONS: Signalling mechanisms associated with perception of the R : FR gradient in canopies were less important than those associated with the irradiance gradient for between-leaf and within-leaf partitioning of photosynthetic resources. The relative importance of the signals differed between species because Carex was the only species for which no indications were found for an involvement of the spectral gradient in perception of canopy density.

Carex Plant↗

Potato aphid, Macrosiphum euphorbiae (Thomas), in tomatoes: plant canopy distribution and binomial sampling on processing tomatoes in California.

A binomial sampling method for the potato aphid, Macrosiphum euphorbiae (Thomas), on processing tomato plants, Lycopersicon esculentum (Mill), is proposed. Relationships between mean number of M. euphorbiae per leaf and proportion of leaves infested [P(I)] with M. euphorbiae for both upper and interior leaves of the processing tomato varieties 'Alta' and 'Halley' are presented. A split-plot design was used with variety, position in the plant canopy, and block as the factors examined through linear regression and analysis of variance (ANOVA). Results supported the hypotheses that M. euphorbiae densities on upper canopy leaves are predictive of densities on inner canopy leaves and that proportion of M. euphorbiae infested leaves are predictive of mean densities per leaf. Mean M. euphorbiae density was greater on 'Alta' than 'Halley' tomato plants, supporting the assumption that 'Alta' is the more susceptible variety. Taylor's Power Law coefficients, a and b, were similar for proportion of M. euphorbiae-infested upper and inner leaves of both 'Alta' and 'Halley'. Taylor's b coefficients ranged from 1.57 to 1.74, indicating a highly clumped distribution for M. euphorbiae.

Animals↗

Sentinel pigeon surveillance for West Nile virus by using lard-can traps at differing elevations and canopy cover classes.

Sentinel pigeons, Columba livia, were installed in lard-can traps at heights of 1.5 m and 7.6-9.1 m within differing canopy cover classes in New York City. Adult mosquitoes were collected weekly from July to October 2002, as were serum samples from each pigeon. Culex pipiens L. and Culex restuans Theobald comprised 97% of mosquitoes collected and were most numerous in canopy-level, forested traps. The West Nile virus (family Flaviviridae, genus Flavivirus, WNV) seroconversion rate was significantly greater for pigeons in canopy-level traps, although seroconversions occurred concurrently with human cases in the city and were of little prognostic value to public health agencies. Our results indicate that sentinel pigeons were most effective for monitoring enzootic transmission of WNV when placed in single-sentinel caging 7.6-9.1 m above ground level.

Animals↗

3D lidar imaging for detecting and understanding plant responses and canopy structure.

Understanding and diagnosing plant responses to stress will benefit greatly from three-dimensional (3D) measurement and analysis of plant properties because plant responses are strongly related to their 3D structures. Light detection and ranging (lidar) has recently emerged as a powerful tool for direct 3D measurement of plant structure. Here the use of 3D lidar imaging to estimate plant properties such as canopy height, canopy structure, carbon stock, and species is demonstrated, and plant growth and shape responses are assessed by reviewing the development of lidar systems and their applications from the leaf level to canopy remote sensing. In addition, the recent creation of accurate 3D lidar images combined with natural colour, chlorophyll fluorescence, photochemical reflectance index, and leaf temperature images is demonstrated, thereby providing information on responses of pigments, photosynthesis, transpiration, stomatal opening, and shape to environmental stresses; these data can be integrated with 3D images of the plants using computer graphics techniques. Future lidar applications that provide more accurate dynamic estimation of various plant properties should improve our understanding of plant responses to stress and of interactions between plants and their environment. Moreover, combining 3D lidar with other passive and active imaging techniques will potentially improve the accuracy of airborne and satellite remote sensing, and make it possible to analyse 3D information on ecophysiological responses and levels of various substances in agricultural and ecological applications and in observations of the global biosphere.

Biosensing Techniques↗