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Cambium

The Cambium investigation is one in a pair of investigations which utilizes the Advanced Biological Research System (ABRS). Cambium seeks definitive evidence that gravity has a direct effect on cambial cells (cells located under the inner bark where secondary growth occurs) in willow, Salix babylonica. The Cambium investigation uses willow plants flown on the International Space Station to better understand the fundamental processes by which plants produce cellulose and lignin, the two main structural materials found in plant matter. On Earth, the nature of wood within tree stems varies depending on position, and that within-tree variation includes differences in cell types and chemistry including lignin and cellulose, two major components of wood influencing wood strength, usefulness and carbon content. Reaction wood is an extreme example of such variation, and it is believed that reaction wood develops as a reaction to the influence of gravity. For the Cambium experiment, young willow plants will be launched to the ISS where their stems will be looped in an attempt to induce reaction wood formation. After on-orbit growth, the plants will be preserved and returned to Earth for analysis. Understanding the role of gravity in wood formation is expected to enable wiser management of forests for carbon sequestration as well as better utilization of trees for wood products. Detailed Research Description: The Cambium experiment will provide an understanding of physiological processes such as gene expression, metabolism and general plant development that are affected in plant systems exposed to space flight. Cambium seeks definitive evidence that gravity has a direct effect on the cambial cells (cells located under the inner bark where secondary growth occurs) that contribute to xylogenesis (reaction wood formation) in willow plants, Salix babylonica. Tension wood fibers differentiate on the upper sides of stems when the stem is altered from its normal (vertical) growth position by looping. This reaction wood response does not occur if the orientation of the plant stem with respect to gravity is not altered. If a localized zone of tension wood should be formed in looped stems under microgravity conditions, this would be the first conclusive evidence that gravity is not required. On the other hand, if a zone of tension wood is not produced in looped stems (subjected to tension on one side, compression on the other) in microgravity, this would be the first definitive evidence that gravity has a direct effect on the cambial cells which contribute to reaction wood formation. Following return to Earth the plants will be analyzed by microscopy and chemical methods.

Savidge, Rodney↗

An evaluation of imaging spectrometry for estimating forest canopy chemistry

High spectral resolution Airborne Imaging Spectrometer (AIS) data were acquired over 20 well-studied Wisconsin forest sites to evaluate the potential of remote sensing for estimating forest canopy chemistry. Intensive nutrient cycling research in these forests demonstrates that canopy lignin content is strongly related to measured annual nitrogen mineralization at the undisturbed sites and may serve as an accurate index for nitrogen cycling rates. Ground measurements were made of foliar biomass and canopy nitrogen and lignin content, the latter within two weeks of the AIS overflight. The spectral data were transformed using derivative techniques modified from laboratroy spectroscopy. Stepwise regression assisted in determining combinations of wavelengths most highly correlated with canopy chemistry and biomass. Strong correlations between AIS data and total canopy lignin content in deciduous forests and canopy lignin concentration (total lignin/biomass) in both deciduous and coniferous stands indicate that imaging spectrometry can be used to estimate canopy lignin content and, from that, the spatial distribution of annual nitrogen mineralization rates.

Wessman, Carol A.↗

High spectral resolution remote sensing of canopy chemistry

Near infrared laboratory spectra have been used for many years to determine nitrogen and lignin concentrations in plant materials. In recent years, similar high spectral resolution visible and infrared data have been available via airborne remote sensing instruments. Using data from NASA's Airborne visible/Infrared Imaging Spectrometer (AVIRIS) we attempt to identify spectral regions correlated with foliar chemistry at the canopy level in temperate forests.

Aber, John D.↗

Application of a two-stream radiative transfer model for leaf lignin and cellulose concentrations from spectral reflectance measurements, part 2

We used the Kubelka-Munk theory of diffuse spectral reflectance in layers to analyze influences of multiple chemical components in leaves. As opposed to empirical approaches to estimation of plant chemistry, the full spectral resolution of laboratory reflectance data was retained in an attempt to estimate lignin or other constituent concentrations from spectral band positions. A leaf water reflectance spectrum was derived from theoretical mixing rules, reflectance observations, and calculations from theory of intrinsic k- and s-functions. Residual reflectance bands were then isolated from spectra of fresh green leaves. These proved hard to interpret for composition in terms of simple two component mixtures such as lignin and cellulose. We next investigated spectral and dilution influences of other possible components (starch, protein). These components, among others, added to cellulose in hypothetical mixtures, produce band displacements similar to lignin, but will disguise by dilution the actual abundance of lignin present in a multicomponent system. This renders interpretation of band positions problematical. Knowledge of end-members and their spectra, and a more elaborate mixture analysis procedure may be called for. Good observational atmospheric and instrumental conditions and knowledge thereof are required for retrieval of expected subtle reflectance variations present in spectra of green vegetation.

Conel, James E.↗

Measurements of canopy chemistry with 1992 AVIRIS data at Blackhawk Island and Harvard Forest

The research described in this paper was designed to determine if high spectral resolution imaging spectrometer data can be used to measure the chemical composition of forest foliage, specifically nitrogen and lignin concentration. Information about the chemical composition of forest canopies can be used to determine nutrient cycling rates and carbon balances in forest ecosystems. This paper will describe the results relating data from the Airborne Visible/Infrared Imaging Spectrometer (AVIRIS) to field measured canopy chemistry at Blackhawk Island, WI and Harvard Forest, MA.

Martin, Mary E.↗

Ecosystem and physiological controls over methane production in northern wetlands

Peat chemistry appears to exert primary control over methane production rates in the Canadian Northern Wetlands Study (NOWES) area. We determined laboratory methane production rate potentials in anaerobic slurries of samples collected from a transect of sites through the NOWES study area. We related methane production rates to indicators of resistance to microbial decay (peat C: N and lignin: N ratios) and experimentally manipulated substrate availability for methanogenesis using ethanol (EtOH) and plant litter. We also determined responses of methane production to pH and temperature. Methane production potentials declined along the gradient of sites from high rates in the coastal fens to low rates in the interior bogs and were generally highest in surface layers. Strong relationships between CH4 production potentials and peat chemistry suggested that methanogenesis was limited by fermentation rates. Methane production at ambient pH responded strongly to substrate additions in the circumneutral fens with narrow lignin: N and C: N ratios (delta CH4/delta EtOH = 0.9-2.3 mg/g) and weakly in the acidic bogs with wide C: N and lignin: N ratios (delta CH4/delta EtOH = -0.04-0.02 mg/g). Observed Q(sub 10) values ranged from 1.7 to 4.7 and generally increased with increasing substrate availability, suggesting that fermentation rates were limiting. Titration experiments generally demonstrated inhibition of methanogenesis by low pH. Our results suggest that the low rates of methane emission observed in interior bogs during NOWES likely resulted from pH and substrate quality limitation of the fermentation step in methane production and thus reflect intrinsically low methane production potentials. Low methane emission rates observed during NOWES will likely be observed in other northern wetland regions with similar vegetation chemistry.

Valentine, David W.↗

Remote sensing of canopy chemistry and nitrogen cycling in temperate forest ecosystems

The use of images acquired by the Airborne Imaging Spectrometer, an experimental high-spectral resolution imaging sensor developed by NASA, to estimate the lignin concentration of whole forest canopies in Wisconsin is reported. The observed strong relationship between canopy lignin concentration and nitrogen availability in seven undisturbed forest ecosystems on Blackhawk Island, Wisconsin, suggests that canopy lignin may serve as an index for site nitrogen status. This predictive relationship presents the opportunity to estimate nitrogen-cycling rates across forested landscapes through remote sensing.

Wessman, Carol A.↗

Vegetation Biochemistry: What Can Imaging Spectrometry Tell Us About Canopies?

Changes in ecosystem processes such as productivity and decomposition may be expressed in the canopy foliar chemistry resulting from altered carbon allocation patterns, metabolic processes and nutrient availability. Understanding carbon balance on land over large regions requires quantitative determination of leaf constituents such as lignin and total nitrogen from remote sensing imaging systems. Results from spectral reflectance measurements of stacked leaves in the laboratory show that spectrum matching techniques are applicable to the derivation of the equivalent liquid water thickness in plants as well as to the extraction of dry leaf matter reflectance spectra from spectra of green leaves. The residual spectra derived by subtracting water spectra from the spectra of green leaves shows a feature at 1.72 micrometers that can be related to the lignin content of the leaves. Oak leaves have a deeper residual absorption feature than do cotton leaves which is consistent with their relative lignin content. Similar results are achieved when deriving the residuals from images taken over areas of grass and pine trees. Imaging spectrometry provides promise in developing images of various foliar biochemical constituents.

Goetz, Alexander F. H.↗

Prediction of leaf chemistry by the use of visible and near infrared reflectance spectroscopy

The chemical content of dry, ground leaf material sampled from deciduous and conifer tree species from sites in Alaska, Wisconsin, and California was estimated using visible and shortwave IR spectroscopy. Seven chemical components - sugar, starch, protein, cellulose, total chlorophyll, lignin, and total nitrogen - were analyzed by wet chemical methods and their concentrations regressed against log 1/rho and first and second differences of log 1/rho (where rho is measured reflectance) at wavelengths selected by stepwise regression. Predictions of chemical concentrations based on cross validation suggest that this technique may be useful for extracting vegetation canopy biochemical information by remote sensing.

Card, Don H.↗

Plant metabolism and cell wall formation in space (microgravity) and on Earth

Variations in cell wall chemistry provide vascular plants with the ability to withstand gravitational forces, as well as providing facile mechanisms for correctional responses to various gravitational stimuli, e.g., in reaction wood formation. A principal focus of our current research is to precisely and systematically dissect the essentially unknown mechanism(s) of vascular plant cell wall assembly, particularly with respect to formation of its phenolic constituents, i.e., lignins and suberins, and how gravity impacts upon these processes. Formation of these phenolic polymers is of particular interest, since it appears that elaboration of their biochemical pathways was essential for successful land adaptation. By extrapolation, we are also greatly intrigued as to how the microgravity environment impacts upon 'normal' cell wall assembly mechanisms/metabolism.

Lewis, Norman G.↗

Estimating key forest ecosystem parameters through remote sensing

Forest canopy chemistry and biomass indicators of ecosystem photosynthesis and decomposition processes are presently studied in view of Airborne Imaging Spectrometer data, which generated spectra from averaged 3 x 3 pixel areas for each of 20 sites for mutual qualitative comparison. Vegetation spectra were strongly differentiated from other cover types by an apparent absorption feature at 1500-1700 nm. Preliminary work with stepwise regression suggests that lignin may play a role in canopy reflectance, and that there is potential for remote detection of forest canopy lignin.

Wessman, Carol A.↗

Seasonal Patterns and Remote Spectral Estimation of Canopy Chemistry Across the Oregon Transect

We examined seasonal changes in canopy chemical concentrations and content in conifer forests growing along a climate gradient in western Oregon, as part of the Oregon Transect Ecosystem Research (OTTER) study. The chemical variables were related to seasonal patterns of growth and production. Statistical comparisons of chemical variables with data collected from two different airborne remote-sensing platforms were also carried out. Total nitrogen (N) concentrations in foliage varied significantly both seasonally and among sites; when expressed as content in the forest canopy, nitrogen varied to a much greater extent and was significantly related to aboveground net primary production (r = 0.99). Chlorophyll and free amino acid concentrations varied more strongly than did total N and may have reflected changes in physiological demands for N. Large variations in starch concentrations were measured from pre- to post-budbreak in all conifer sites. Examination of remote-sensing data from two different airborne instruments suggests the potential for remote measurement of some canopy chemicals. Multivariate analysis of high-resolution spectral data in the near infrared region indicated significant correlations between spectral signals and N concentration and canopy N content; the correlation with canopy N content was stronger and was probably associated in part with water absorption features of the forest canopy. The spectral bands that were significantly correlated with lignin concentration and content were similar to bands selected in the other laboratory and airborne studies; starch concentrations were not significantly related to spectral reflectance data. Strong relationships between the spectral position of specific reflectance features in the visible region and chlorophyll were also found.

Matson, Pamela↗

Gravity as a biochemical determinant

The existence of obvious morphological and physiological changes in living systems exposed to altered gravity immediately informs us that prior changes have taken place in the chemistry of exposed cells, tissues and organs. These changes include transients that return more or less promptly to the norm when the system is restored to the terrestrial g-field. For example, altered serum hormone and electrolyte levels in man, which appear to reflect successful adaptation to the conditions of orbital weightlessness, disappear shortly after return to Earth. Other changes--in mineral and protein constituents of the skeletal system in man, and cell wall composition in plants--are more persistent or even permanent. Hypogravitational departures from the norm include not only "weightlessness" as achieved in orbit, but also experimental modes of compensation, on the clinostat or by flotation. These techniques are useful in the study of hypogravity but cannot replace fully the weightless environment. Plant ethylene and peroxidase both increase under orbital, clinostat and/or flotation conditions whereas 3-phosphoglyceraldehyde-dehydrogenase increases under orbital but not clinostat conditions; cytochrome reductase and malic dehydrogenase levels are affected by the clinostat, but not by actual weightless conditions. How do the altered organismal biochemistries induced by the centrifuge and the clinostat relate to one another? Does gravity operate on living systems as a continuous variable from 0 to superterrestrial values, or do deviations from g(earth) generate non-uniform, discontinuous stress responses, irrespective of sign? In plants, measurements of wall lignin content and peroxidase activity yield opposite answers. Given the limited data so far available we will consider the meaning of these contradictions.

Review↗