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Phytochrome-mediated responses: Implications for controlled environment research facilities

Light is undoubtedly the most important environmental variable for plant growth and development; plants not only use radiant energy in photosynthesis, they also respond to the quantity, quality, direction and timing of incident radiation through photomorphogenic response that can have huge effects on the rate of growth and the pattern of development. It is surprising, therefore, that the manufacturers and suppliers of controlled environment facilities have been singularly uninventive in the design of the lighting assemblies they provide. The consumer has one choice only - a lighting assembly that provides irradiance levels usually only a fraction of sunlight, and a control system that is limited to regulating the timing of the on-off switch. The reasons for these limitations are partly technological, but in the main they result from ignorance on the part of both the consumer and the manufacturer. A specific and powerful example of this ignorance relates to the importance of the so-called far-red wavelengths (FR = 700-800 nm). Because the human eye can hardly detect wavelengths above 700 nm, and photosynthesis also cuts off at about 700 nm, the majority of plant and crop physiologists are still almost completely unaware that FR radiation can have massive effects on growth rate and development. In consequence, most growth cabinets have light sources based on fluorescent tubes, and provide very little FR apart from that emitted by a token number of small incandescent bulbs. Larger growth facilities often use broader spectrum light sources, but growth facilities that provide the capability to vary the FR incident upon the plants are about as abundant as seals in the Sahara. This article sets the background of the significance of FR radiation in the natural environment and its importance for plant growth and development in the hope that it might inform intelligently those concerned with improving the design of plant growth facilities.

Smith, Harry↗

Spectral filtering for plant production

Both plants and animals have one general commonality in their perception of light. They both are sensitive primarily to the 400 to 700 nm wavelength portion of the electromagnetic spectrum. This is referred to as the visible spectrum for animals and as the photosynthetically active radiation (PAR) spectrum for plants. Within this portion of the spectrum, animals perceive colors. Relatively recently it has been learned that within this same spectral range plants also demonstrate varying responses at different wavelengths, somewhat analogous to the definition of various colors at specific wavelengths. Although invisible to the human eye, portions of the electromagnetic spectrum on either side of the visible range are relatively inactive photosynthetically but have been found to influence important biological functions. These portions include the ultraviolet (UV approximately equal to 280-400 nm) and the far-red (FR approximately equal to 700-800 nm). The basic photoreceptor of plants for photosynthesis is chlorophyll. It serves to capture radiant energy which combined with carbon dioxide and water produces oxygen and assimulated carbon, used for the synthesis of cell wall polysaccarides, proteins, membrane lipids and other cellular constituents. The energy and carbon building blocks of photosynthesis sustain growth of plants. On the other hand, however, there are other photoreceptors, or pigments, that function as signal transducers to provide information that controls many physiological and morphological responses of how a plant grows. Known photomorphogenic receptors include phytochrome (the red/far-red sensor in the narrow bands of 655-665 nm and 725-735 nm ranges, respectively) and 'cryptochrome' (the hypothetical UV-B sensor in the 280-320 nm range). Since the USDA team of W. L. Butler, S. B. Hendricks, H. A. Borthwick, H. A. Siegleman and K. Norris in Beltsville, MD detected by spectroscopy, extracted and identified phytochrome as a protein in the 1950's, many other investigators have found evidence of its control functions in plants. Considerably less, however, is known about the yet non-isolated cryptochrome. The information-transferring roles of photoreceptors in plants at specific spectral ranges quite naturally stimulated plant scientists and engineers to consider physically manipulating light to achieve desired physiological and morphological characteristics. One way to manipulate light is to filter it through materials that selectively transmit portions of the sun's spectrum in and near the PAR range.

Young, Roy E.↗

Modelling Temporal Variability in the Carbon Balance of a Spruce/Moss Boreal Forest

A model of the daily carbon balance of a black spruce/feathermoss boreal forest ecosystem was developed and results compared to preliminary data from the 1994 BOREAS field campaign in northern Manitoba, Canada. The model, driven by daily weather conditions, simulated daily soil climate status (temperature and moisture profiles), spruce photosynthesis and respiration, moss photosynthesis and respiration, and litter decomposition. Model agreement with preliminary field data was good for net ecosystem exchange (NEE), capturing both the asymmetrical seasonality and short-term variability. During the growing season simulated daily NEE ranged from -4 g C m(exp -2) d(exp -1) (carbon uptake by ecosystem) to + 2 g C m(exp -2) d(exp -1) (carbon flux to atmosphere), with fluctuations from day to day. In the early winter simulated NEE values were + 0.5 g C m(exp -2) d(exp -1), dropping to + 0.2 g C m(exp -2) d(exp -1) in mid-winter. Simulated soil respiration during the growing season (+ 1 to + 5 g C m(exp -2) d(exp -1)) was dominated by metabolic respiration of the live moss, with litter decomposition usually contributing less than 30% and live spruce root respiration less than 10% of the total. Both spruce and moss net primary productivity (NPP) rates were higher in early summer than late summer. Simulated annual NEE for 1994 was -51 g C m(exp -2) y(exp -1), with 83% going into tree growth and 17% into the soil carbon accumulation. Moss NPP (58 g C m(exp -2) d(exp -1)) was considered to be litter (i.e. soil carbon input; no net increase in live moss biomass). Ecosystem respiration during the snow-covered season (84 g Cm(exp -2)) was 58% of the growing season net carbon uptake. A simulation of the same site for 1968-1989 showed about 10-20% year-to-year variability in heterotrophic respiration (mean of + 113 g C m-2 y@1). Moss NPP ranged from 19 to 114 g C m(exp -2) y(exp -1); spruce NPP from 81 to 150 g C nt-2 y,@l; spruce growth (NPP minus litterfall) from 34 to 103 g C m(exp -2) y(exp -1); NEE ranged from +37 to -142 g C m(exp -2) y(exp -1). Values for these carbon balance terms in 1994 were slightly smaller than the 1969 - 89 means. Higher ecosystem productivity years (more negative NEE) generally had early springs and relatively wet summers; lower productivity years had late springs and relatively dry summers.

Frolking, S.↗

Use of Molecular Fossils for the Interpretation of Paleoenvironments on Early Earth: The Synthesis of Lipid Biomarkers by Methane-Oxidizing Bacteria

Over the course of Earth's history, the most important biological influence has come from the evolution of oxygenic photosynthesis and the development of an oxygen-rich biosphere. Although the availability of free oxygen had profound effects on subsequent biological and planetary evolution, clear paleobiological evidence of the timing of this transition is lacking. Recent technical advances in the microanalysis of organic matter has made it possible to detect residual molecules (biomarkers) in proterozoic (2.5 to 0.6 billion years) sedimentary rock characteristic of specific groups of microorganisms. When coupled with the carbon isotopic fractionations characteristic of biological systems and the new field of compound specific isotope analysis, biomarkers could prove to be a powerful tool for decoding ancient biochemistry from the geological record. We have been studying the carbon isotope fractionations associated with the synthesis of organic biomarker molecules in several types of methane-oxidizing bacteria that should have been key to carbon cycling in paleoenvironments after the evolution of oxygenic photosynthesis.

Jahnke, Linda L.↗

Measurement of Effective Canopy Temperature: The Missing Link to Modeling Transpiration in Controlled Environments

The objectives were to apply energy balance principles to plant canopies, and to determine which parameters are essential for predicting plant canopy transpiration (E) in controlled environments. Transpiration was accurately measured in a gas-exchange system. Absorbed radiation (R(sub abs)) by the canopy was measured with a net radiometer and calculated from short and long-wave radiation components. Average canopy foliar temperature T(sub L) can be measured with an infrared radiometer, but since T(sub L) is seldom uniform, a weighed average measurement of T(sub L) must be made. The effective canopy temperature T(sub C) is that temperature that balances the energy flux between absorbed radiation and latent heat L(sub E) and sensible heat (H) fluxes. TC should exactly equal air temperature T(sub A) when L(sub E) equals R(sub abs). When unnecessary thermal radiation from the lighting system is removed by a water filter, the magnitude of L(sub E) from transpiration approaches Rabs and T(sub C) is close to T(sub A). Unlike field models, we included the energy used in photosynthesis and found that up to 10% of Rabs was used in photosynthesis. We calculated aerodynamic conductance for H from measurements of wind speed and canopy height using the wind profile equation. Canopy aerodynamic conductance ranged from.03 to.04 m/s for wind speeds from.6 to 1 m/s; thus a 0.1 C canopy to air temperature difference results in a sensible heat flux of about 4 W/sq m, which is only 1% of R(sub abs). We examined the ability of wide angle infrared transducers to accurately integrate T(sub L) from the top to the bottom of the canopy. We measured evaporation from the hydroponic media to be approximately 1 micro mol/sq m s or 10% of R(sub abs). This result indicates that separating evaporation from transpiration is more important than exact measurement of canopy temperature.

Monje, O. A.↗

Biogenic Methane and the Rise of Oxygen

Oxygenic photosynthesis does not make the rise of oxygen inevitable. What is required is that reductant and oxygen be separated and permanently segregated. The usual picture for Earth is that oxygenic photosynthesis split CO2 into carbon and oxygen, with the carbon buried in sediments and the oxygen mostly taken up by oxides of iron and sulfur. The relatively small atmospheric reservoir of O2 is regulated by the carbon burial rate, reaction with volcanic and metamorphic gases, and oxidation of reduced carbon released as old sediments weather. Absent from this picture is a distinction between the Archean and modern times: on average, carbon burial fluxes would have been matched by oxygen losses then as now. Separation of reductant from oxidant is only provisional. No net oxidation of the continents occurs, and so no change of diagenetic, metamorphic, or volcanic gases is expected. Nor would any change in oxidative weathering be expected. Something more than carbon burial is required to make the Archean different. The escape of hydrogen to space permanently separates the reductant from the oxidant. Hydrogen escape is widely believed to have led to the present highly oxidized states of Mars and Venus. Hydrogen escape has usually been thought small for Archaean Earth, because water vapor is cold-trapped at the troposphere and thus held to levels of a few ppmv in the stratosphere. This cold trapping renders hydrogen escape negligible. However, methane is not cold trapped, and its expected abundance in the Archaean, given low oxygen levels and a biogenic source, would have been high, probably more than 100 times present. At such levels methane would have driven geologically significant levels of hydrogen escape. Additional information is contained in the original extended abstract.

Catling, David↗

Chloroplast Ultrastructure of the Alga Phaeocystis antarctica Karsten: A New Structural Model Using Electron Tomography

Understanding the light-harvesting properties of algae and higher plants are a fundamental topic in photosynthesis research. Using thick sections obtained from fixed and embedded cultures of colonial P antarctica, we calculate tomographic reconstructions of individual chloroplasts under light-limiting and saturating conditions for net photosynthesis. Our goal is to gain an understanding of the continuity of thylakoid membranes and understand the spatial relationship between the pyrenoid, the starch containing organelle, and thylakoid membranes. We found that Phaeocystis showed considerable morphological and physiological flexibility in response to environmental light levels. We found that the thylakoids generally run parallel to the chloroplast membrane with many junctures and bifurcations, many of which are in contact with the chloroplast membrane itself. The considerable flexibility in the. thylakoid membranes allows for the accommodation of the pyrenoid structure. The arrangement of the thylakoids within these structures resemble those found in new structures of mitochondria cristae. We present a new structural model for algal chloroplasts which greatly revises current concepts of thylakoid membrane structure in relation to photoacclimation.

Moisan, Tiffany A.↗

Benthic Marine Cyanobacterial Mat Ecosystems: Biogeochemistry and Biomarkers

Cyanobacterial mats are complete ecosystems that can include processes of primary production, diagenesis and lithification. Light sustains oxygenic photosynthesis, which in turn provides energy, organic matter and oxygen to the community. Due to both absorption and scattering phenomena, incident light is transformed with depth in the mat, both in intensity and spectral composition. Mobile photo synthesizers optimize their position with respect to this light gradient. When photosynthesis ceases at night, the upper layers of the mat become reduced and sulfidic. Counteracting gradients of oxygen and sulfide combine to provide daily-contrasting environments separated on a scale of a few mm. The functional complexity of mats, coupled with the highly proximal and ordered spatial arrangement of biota, offers the potential for a staggering number of interactions. At a minimum, the products of each functional group of microorganisms affect the other groups both positively and negatively. For example, cyanobacteria generate organic matter (potential substrates) but also oxygen (a toxin for many anaerobes). Anaerobic activity recycles nutrients to the photosynthesizers but also generates potentially toxic sulfide. The combination of benefits and hazards of light, oxygen and sulfide promotes the allocation of the various essential mat processes between light and dark periods, and to various depths in the mat. Observations of mats have produced numerous surprises. For example, obligately anaerobic processes can occur in the presence of abundant oxygen, highly reduced gases are produced in the presence of abundant sulfate, meiofauna thrive at high sulfide concentrations, and the mats' constituent populations respond to environmental changes in complex ways. While photosynthetic bacteria dominate the biomass and productivity of the mat, nonphotosynthetic, anaerobic processes constitute the ultimate biological filter on the ecosystem's emergent biosignatures, including those sedimentary textures, organic compounds, and minerals that enter the fossil record. The ability of cyanobacterial mats to channel abundant solar energy into the creation and maintenance of complex structures and processes has created a multitude of consequences, both for sedimentation and for the early evolution of our biosphere.

DesMarais, David J.↗

Estimating Net Primary Productivity Using Satellite and Ancillary Data

The net primary productivity (C) or the annual rate of carbon accumulation per unit ground area by terrestrial plant communities is the difference of gross photosynthesis (A(sub g)) and respiration (R) per unit ground area. Available field observations show that R is a large and variable fraction of A(sub g), although it is generally recognized that there are considerable difficulties in determining these fluxes, and thus pose challenge in assessing the accuracy. Further uncertainties arise in extrapolating field measurements (which are acquired over a hectare or so area) to regional scale. Here, an approach is presented for determining these fluxes using satellite and ancillary data to be representative of regional scale and allow assessment of interannual variation. A, has been expressed as the product of radiation use efficiency for gross photosynthesis by an unstressed canopy and intercepted photosynthetically active radiation, which is then adjusted for stresses due to soil water shortage and temperature away from optimum. R has been calculated as the sum of growth and maintenance components (respectively, R(sub g) and R(sub m)).The R(sub m) has been determined from nitrogen content of plant tissue per unit ground area, while R(sub g) has been obtained as a fraction of the difference of A(sub g) and R(sub m). Results for five consecutive years (1986-1990) are presented for the Amazon-Tocontins, Mississippi, and Ob River basins.

Choudhury, Bhaskar J.↗

Controls on Variations of Surface Energy, Water, and Carbon Budgets within Large-Scale Amazon Basin

A key research focus of the LBA Research Program is understanding the space-time variations in interlinked surface energy, water, and carbon budgets, the controls on these variations, and the implications of these controls on the carbon sequestering capacity of the large scale forest-pasture system that dominates the Amaz6nia landscape. Quantification of these variations and controls are investigated by a combination of in situ measurements, remotely sensed measurements from space, and a realistically forced hydrometeorological model coupled to a carbon assimilation model, capable of simulating details within the surface energy and water budgets along with the principle processes of photosynthesis and respiration. Herein we describe the results of an investigation concerning the space-time controls of carbon sources and sinks distributed over the large scale Amazon basin. The results are derived from a carbon-water-energy budget retrieval system for the large scale Amazon basin, which uses a coupled carbon assimilation-hydrometeorological model as an integrating system, forced by both in situ meteorological measurements and remotely sensed radiation and precipitation fluxes obtained from a combination of GOES, SSM/I, TOMS, and TRh4M satellite measurements. Results include validation of (a) retrieved surface radiation and precipitation fluxes based on 30-min averaged surface measurements taken at Ji-Parani in Rondania and Manaus in Amazonas, and (b) modeled sensible, latent, and C02 fluxes based on tower measurements taken at Reserva Jaru, Manaus and Fazenda Nossa Senhora. The space-time controls on carbon sequestration are partitioned into sets of factors classified by: (1) above canopy meteorology, (2) incoming surface radiation, (3) precipitation interception, and (4) indigenous stomatal processes varied over the different land covers of pristine rainforest, partially, and fully logged rainforests, and pasture lands. These are the principle meteorological, thermodynamical, hydrological, and biophysical control paths which perturb net carbon fluxes and sequestration, produce time-space switching of carbon sources and sinks, undergo modulation through atmospheric boundary layer feedbacks, and respond to any discontinuous intervention on the landscape itself such as produced by human intervention in converting rainforest to pasture or conducting selective/clearcut logging operations. The results demonstrate how relative carbon sequestration capacity of the Amazonian ecosystem responds to these controls, and how interpretation of space-time heterogeneities in carbon sequestration depends on a fairly exact quantification of the interacting non-linear properties of photosynthesis in response to incoming solar flux, air-canopy temperatures, and leaf water interception -- and soil respiration in response to upper layer soil temperature and water content. The results also show how the interpretation of the control processes is highly sensitive to the scales at which the surface fluxes are analyzed.

Smith, Eric A.↗

Sulfur Cycling Mediates Calcium Carbonate Geochemistry in Modern Marine Stromatolites

Modem marine stromatolites forming in Highborne Cay, Exumas (Bahamas), contain microbial mats dominated by Schizothrix. Although saturating concentrations of Ca2+ and CO32- exist, microbes mediate CaCO3 precipitation. Cyanobacterial photosynthesis in these stromatolites aids calcium carbonate precipitation by removal of HS+ through CO2 use. Photorespiration and exopolymer production predominantly by oxygenic phototrophs fuel heterotrophic activity: aerobic respiration (approximately 60 umol/sq cm.h) and sulfate reduction (SR; 1.2 umol SO42-/sq cm.h) are the dominant C- consuming processes. Aerobic microbial respiration and the combination of SR and H2S oxidation both facilitate CaCO3 dissolution through H+ production. Aerobic respiration consumes much more C on an hourly basis, but duel fluctuating O2 and H2 depth profiles indicate that overall, SR consumes only slightly less (0.2-0.5) of the primary production. Moreover, due to low O2 concentrations when SR rates are peaking, reoxidation of the H2S formed is incomplete: both thiosulfate and polythionates are formed. The process of complete H2S oxidation yields H+. However, due to a low O2 concentration late in the day and relatively high O2 concentrations early in the following morning, a two-stage oxidation takes place: first, polythionates are formed from H2S, creating alkalinity which coincides with CaCO3 precipitation; secondly, oxidation of polythionates to sulfate yields acidity, resulting in dissolution, etc. Vertical profiles confirmed that the pH peaked late in the afternoon (greater than 8.8) and had the lowest values (less than 7.4) early in the morning. Thus, the effect of this S-cycling through alkalinity production, followed by acidification during H2S oxidation, results in a six times stronger fluctuation in acidity than photosynthesis plus aerobic respiration accomplish. This implies that anaerobic processes play a pivotal role in stromatolite formation.

Visscher, P. T.↗

Quantitative Relationships between Photosynthetic, Nitrogen Fixing, and Fermentative H2 Metabolism in a Photosynthetic Microbial Mat

The ultimate potential of any microbial ecosystem to contribute chemically to its environment - and therefore, to impact planetary biogeochemistry or to generate recognizable biosignatures - depends not only on the individual metabolic capabilities of constituent organisms, but also on how those capabilities are expressed through interactions with neighboring organisms. This is particularly important for microbial mats, which compress an extremely broad range of metabolic potential into a small and dynamic system. H2 participates in many of these metabolic processes, including the major elemental cycling processes of photosynthesis, nitrogen fixation, sulfate reduction, and fermentation, and may therefore serve as a mediator of microbial interactions within the mat system. Collectively, the requirements of energy, electron transfer, and biomass element stoichiometry suggest quantitative relationships among the major element cycling processes, as regards H2 metabolism We determined experimentally the major contributions to 32 cycling in hypersaline microbial mats from Baja California, Mexico, and compared them to predicted relationships. Fermentation under dark, anoxic conditions is quantitatively the most important mechanism of H2 production, consistent with expectations for non-heterocystous mats such as those under study. Up to 16% of reducing equivalents fixed by photosynthesis during the day may be released by this mechanism. The direct contribution of nitrogen fixation to H2 production is small in comparison, but this process may indirectly stimulate substantial H2 generation, by requiring higher rates of fermentation. Sulfate reduction, aerobic consumption, diffusive and ebulitive loss, and possibly H2-based photoreduction of CO2 serve as the principal H2 sinks. Collectively, these processes interact to create an orders-of-magnitude daily variation in H2 concentrations and fluxes, and thereby in the oxidation-reduction potential that is imposed on microbial processes occuring within the mat matrix.

Hoehler, Tori M.↗

Earth's early biosphere

Understanding our own early biosphere is essential to our search for life elsewhere, because life arose on Earth very early and rocky planets shared similar early histories. The biosphere arose before 3.8 Ga ago, was exclusively unicellular and was dominated by hyperthermophiles that utilized chemical sources of energy and employed a range of metabolic pathways for CO2 assimilation. Photosynthesis also arose very early. Oxygenic photosynthesis arose later but still prior to 2.7 Ga. The transition toward the modern global environment was paced by a decline in volcanic and hydrothermal activity. These developments allowed atmospheric O2 levels to increase. The O2 increase created new niches for aerobic life, most notably the more advanced Eukarya that eventually spawned the megascopic fauna and flora of our modern biosphere.

Review↗

Accuracy of quantum sensors measuring yield photon flux and photosynthetic photon flux

Photosynthesis is fundamentally driven by photon flux rather than energy flux, but not all absorbed photons yield equal amounts of photosynthesis. Thus, two measures of photosynthetically active radiation have emerged: photosynthetic photon flux (PPF), which values all photons from 400 to 700 nm equally, and yield photon flux (YPF), which weights photons in the range from 360 to 760 nm according to plant photosynthetic response. We selected seven common radiation sources and measured YPF and PPF from each source with a spectroradiometer. We then compared these measurements with measurements from three quantum sensors designed to measure YPF, and from six quantum sensors designed to measure PPF. There were few differences among sensors within a group (usually <5%), but YPF values from sensors were consistently lower (3% to 20%) than YPF values calculated from spectroradiometric measurements. Quantum sensor measurements of PPF also were consistently lower than PPF values calculated from spectroradiometric measurements, but the differences were <7% for all sources, except red-light-emitting diodes. The sensors were most accurate for broad-band sources and least accurate for narrow-band sources. According to spectroradiometric measurements, YPF sensors were significantly less accurate (>9% difference) than PPF sensors under metal halide, high-pressure sodium, and low-pressure sodium lamps. Both sensor types were inaccurate (>18% error) under red-light-emitting diodes. Because both YPF and PPF sensors are imperfect integrators, and because spectroradiometers can measure photosynthetically active radiation much more accurately, researchers should consider developing calibration factors from spectroradiometric data for some specific radiation sources to improve the accuracy of integrating sensors.

NASA Center KSC↗

Photomorphogenesis and photoassimilation in soybean and sorghum grown under broad spectrum or blue-deficient light sources

The role of blue light in plant growth and development was investigated in soybean (Glycine max [L.] Merr. cv Williams) and sorghum (Sorghum bicolor [L.] Moench. cv Rio) grown under equal photosynthetic photon fluxes (approximately 500 micromoles per square meter per second) from broad spectrum daylight fluorescent or blue-deficient, narrow-band (589 nanometers) low pressure sodium (LPS) lamps. Between 14 and 18 days after sowing, it was possible to relate adaptations in photosynthesis and leaf growth to dry matter accumulation. Soybean development under LPS light was similar in several respects to that of shaded plants, consistent with an important role for blue light photoreceptors in regulation of growth response to irradiance. Thus, soybeans from LPS conditions partitioned relatively more growth to leaves and maintained higher average leaf area ratios (mean LAR) that compensated lower net assimilation rates (mean NAR). Relative growth rates were therefore comparable to plants from daylight fluorescent lamps. Reductions in mean NAR were matched by lower rates of net photosynthesis (A) on an area basis in the major photosynthetic source (first trifoliolate) leaf. Lower A in soybean resulted from reduced leaf dry matter per unit leaf area, but lower A under LPS conditions in sorghum correlated with leaf chlorosis and reduced total nitrogen (not observed in soybean). In spite of a lower A, mean NAR was larger in sorghum from LPS conditions, resulting in significantly greater relative growth rates (mean LAR was approximately equal for both light conditions). Leaf starch accumulation rate was higher for both species and starch content at the end of the dark period was elevated two- and three-fold for sorghum and soybean, respectively, under LPS conditions. Possible relations between starch accumulation, leaf export, and plant growth in response to spectral quality were considered.

NASA Discipline Life Support Systems↗

Photosynthetic action spectra and adaptation to spectral light distribution in a benthic cyanobacterial mat

We studied adaptation to spectral light distribution in undisturbed benthic communities of cyanobacterial mats growing in hypersaline ponds at Guerrero Negro, Baja California, Mexico. Microscale measurements of oxygen photosynthesis and action spectra were performed with microelectrodes; spectral radiance was measured with fiber-optic microprobes. The spatial resolution of all measurements was 0.1 mm, and the spectral resolution was 10 to 15 nm. Light attenuation spectra showed absorption predominantly by chlorophyll a (Chl a) (430 and 670 nm), phycocyanin (620 nm), and carotenoids (440 to 500 nm). Blue light (450 nm) was attenuated 10-fold more strongly than red light (600 nm). The action spectra of the surface film of diatoms accordingly showed activity over the whole spectrum, with maxima for Chl a and carotenoids. The underlying dense Microcoleus population showed almost exclusively activity dependent upon light harvesting by phycobilins at 550 to 660 nm. Maximum activity was at 580 and 650 nm, indicating absorption by phycoerythrin and phycocyanin as well as by allophycocyanin. Very little Chl a-dependent activity could be detected in the cyanobacterial action spectrum, even with additional 600-nm light to excite photosystem II. The depth distribution of photosynthesis showed detectable activity down to a depth of 0.8 to 2.5 mm, where the downwelling radiant flux at 600 nm was reduced to 0.2 to 0.6% of the surface flux.

NASA Discipline Exobiology↗

Long-distance signaling within Coleus x hybridus leaves; mediated by changes in intra-leaf CO2?

Rapid long-distance signaling in plants can occur via several mechanisms, including symplastic electric coupling and pressure waves. We show here in variegated Coleus leaves a rapid propagation of electrical signals that appears to be caused by changes in intra-leaf CO2 concentrations. Green leaf cells, when illuminated, undergo a rapid depolarization of their membrane potential (Vm) and an increase in their apoplastic pH (pHa) by a process that requires photosynthesis. This is followed by a slower hyperpolarization of Vm and apoplastic acidification, which do not require photosynthesis. White (chlorophyll-lacking) leaf cells, when in isolated white leaf segments, show only the slow response, but when in mixed (i.e. green and white) segments, the rapid Vm depolarization and increase in pHa propagate over more than 10 mm from the green to the white cells. Similarly, these responses propagate 12-20 mm from illuminated to unilluminated green cells. The fact that the propagation of these responses is eliminated when the leaf air spaces are infiltrated with solution indicates that the signal moves in the apoplast rather than the symplast. A depolarization of the mesophyll cells is induced in the dark by a decrease in apoplastic CO2 but not by an increase in pHa. These results support the hypothesis that the propagating signal for the depolarization of the white mesophyll cells is a photosynthetically induced decrease in the CO2 level of the air spaces throughout the leaf.

NASA Discipline Plant Biology↗

Radiation: microbial evolution, ecology, and relevance to mars missions

Ultraviolet (UV) radiation has been an important environmental parameter during the evolution of life on Earth, both in its role as a mutagen and as a selective agent. This was probably especially true during the time from 3.8 to 2.5 billion years ago, when atmospheric ozone levels were less than 1% of present levels. Early Mars may not have had an "ozone shield" either, and it never developed a significant one. Even though Mars is farther away from the Sun than the Earth, a substantial surficial UV flux is present on Mars today. But organisms respond to dose rate, and on Mars, like on Earth, organisms would be exposed to diurnal variations in UV flux. Here we present data on the effect of diurnal patterns of UV flux on microbial ecosystems in nature, with an emphasis on photosynthesis and DNA synthesis effects. These results indicate that diurnal patterns of metabolism occur in nature with a dip in photosynthesis and DNA synthesis in the afternoon, in part regulated by UV flux. Thus, diurnal patterns must be studied in order to understand the effect of UV radiation in nature. The results of this work are significant to the success of human missions to Mars for several reasons. For example, human missions must include photosynthetic organisms for food production and likely oxygen production. An evolutionary approach suggests which organisms might be best suited for high UV fluxes. The diurnal aspect of these studies is critical. Terraforming is a potential goal of Mars exploration, and it will require studies of the effect of Martian UV fluxes, including their diurnal changes, on terrestrial organisms. Such studies may suggest that diurnal changes in UV only require mitigation at some times of day or year.

Review, Tutorial↗