Characterizing the purple Earth: Modelling the globally-integrated spectral variability of the Archean Earth (The Astrophysical Journal, 2013)

The ongoing searches for exoplanetary systems have revealed a wealth of planets with diverse physical properties. Planets even smaller than the Earth have already been detected, and the efforts of future missions are placed on the discovery, and perhaps characterization, of small rocky exoplanets within the habitable zone of their stars. Clearly what we know about our planet will be our guideline for the characterization of such planets. But the Earth has been inhabited for at least 3.8 Ga, and its appearance has changed with time. Here, we have studied the Earth during the Archean eon, 3.0 Ga ago. At that time one of the more widespread life forms on the planet were purple bacteria. These bacteria are photosynthetic microorganisms and can inhabit both aquatic and terrestrial environments. Here, we used a radiative transfer model to simulate the visible and near-IR radiation reflected by our planet, taking into account several scenarios regarding the possible distribution of purple bacteria over continents and oceans. We find that purple bacteria have a reflectance spectrum which has a strong reflectivity increase, similar to the red edge of leafy plants, although shifted redwards. This feature produces a detectable signal in the disk-averaged spectra of our planet, depending on cloud amount and on purple bacteria concentration/distribution. We conclude that by using multi-color photometric observations, it is possible to distinguish between an Archean Earth in which purple bacteria inhabit vast extensions of the planet, and a present-day Earth with continents covered by deserts, vegetation or microbial mats.

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Photosystem trap energies and spectrally-dependent energy-storage efficiencies in the Chl d-utilizing cyanobacterium, Acaryochloris marina (BBA Bioenergetics, 2013)

Acaryochloris marina is the only species known to utilize chlorophyll (Chl) d as a principal photopigment. The peak absorption wavelength of Chl d is redshifted ≈40nm in vivo relative to Chl a, enabling this cyanobacterium to perform oxygenic phototrophy in niche environments enhanced in far-red light. We present measurements of the in vivo energy-storage (E-S) efficiency of photosynthesis in A. marina, obtained using pulsed photoacoustics (PA) over a 90-nm range of excitation wavelengths in the red and far-red. Together with modeling results, these measurements provide the first direct observation of the trap energies of PSI and PSII, and also the photosystem-specific contributions to the total E-S efficiency. We find the maximum observed efficiency in A. marina (40±1% at 735nm) is higher than in the Chl a cyanobacterium Synechococcus leopoliensis (35±1% at 690nm). The efficiency at peak absorption wavelength is also higher in A. marina (36±1% at 710nm vs. 31±1% at 670nm). In both species, the trap efficiencies are ≈40% (PSI) and ≈30% (PSII). The PSI trap in A. marina is found to lie at 740±5nm, in agreement with the value inferred from spectroscopic methods. The best fit of the model to the PA data identifies the PSII trap at 723±3nm, supporting the view that the primary electron-donor is Chl d, probably at the accessory (Chl(D1)) site. A decrease in efficiency beyond the trap wavelength, consistent with uphill energy transfer, is clearly observed and fit by the model. These results demonstrate that the E-S efficiency in A. marina is not thermodynamically limited, suggesting that oxygenic photosynthesis is viable in even redder light environments.

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Efficiency of photosynthesis in a Chl d-utilizing cyanobacterium is comparable to or higher than that in Chl a-utilizing oxygenic species (Biochimica Et Biophysica Acta-Bioenergetics, 2011)

The cyanobacterium Acaryochloris marina uses chlorophyll d to carry out oxygenic photosynthesis in environments depleted in visible and enhanced in lower-energy, far-red light. However, the extent to which low photon energies limit the efficiency of oxygenic photochemistry in A. marina is not known. Here, we report the first direct measurements of the energy-storage efficiency of the photosynthetic light reactions in A. marina whole cells, and find it is comparable to or higher than that in typical, chlorophyll a-utilizing oxygenic species. This finding indicates that oxygenic photosynthesis is not fundamentally limited at the photon energies employed by A. marina, and therefore is potentially viable in even longer-wavelength light environments.

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A clumped-foliage canopy radiative transfer model for a Global Dynamic Terrestrial Ecosystem Model II: Comparison to measurements (Agricultural and Forest Meteorology, 2010)

In a previous paper, we developed an analytical clumped two-stream model (ACTS) of canopy radiative transfer from an analytical geometric-optical and radiative transfer (GORT) scheme (Ni-Meister et al., 2010). The ACTS model accounts for clumping of foliage and the influence of trunks in vegetation canopies for modeling of photosynthesis, radiative fluxes and surface albedo in dynamic global vegetation models (DGVMs), and particularly for the Ent Dynamic Global Terrestrial Ecosystem Model (DGTEM). This study evaluates the gap probability and transmittance estimates from the ACTS model by comparing the modeled results with ground-based data, as well as with the original full GORT model and a layered Beer’s law scheme. The ground data used in this study include vertical profile measurements of incident photosynthetically active radiation (PAR) in (1) mixed deciduous forests in Morgan-Monroe State Forest, IN, USA, (2) coniferous forests in central Canada, (3) mixed deciduous forests in Harvard Forest, MA, and (4) ground lidar measurements of the canopy gap fraction in woodland in Australia.

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A clumped-foliage canopy radiative transfer model for a global dynamic terrestrial ecosystem model. I: Theory (Agricultural and Forest Meteorology, 2010)

This study develops a simple but physically based canopy radiative transfer scheme for photosynthesis, radiative fluxes and surface albedo estimates in dynamic global vegetation models (DGVMs), and particularly for the Ent Dynamic Global Terrestrial Ecosystem Model (Ent DGTEM). The Ent DGTEM can represent vegetation in mixed as opposed to homogeneous canopies. With active growth and competition, it must predict radiative transfer for dynamically changing vegetation structure, and requires computational speed for coupling with atmospheric general circulation models (GCMs). The canopy radiative transfer scheme accounts for both vertical and horizontal heterogeneity of plant canopies by combining the simple two-stream scheme with a well-described actual vertical foliage profile, an analytically derived foliage clumping factor from geometric optical theory, and, for needleleaf trees, an empirical needle-to-shoot-clumping factor. In addition, the model accounts for the effect of trunks, which is significant in bare canopies. This model provides better radiation estimates (light profiles, albedo) than the two-stream scheme currently being used in most GCMs to describe light interactions with vegetation canopies. This scheme has the same computational cost as the current typical scheme being used in GCMs, but promises to provide better canopy radiative transfer estimates for DGVMs, particularly those that model heterogeneous vegetation canopies.

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Spectral signatures of photosynthesis. I. Review of Earth organisms (Astrobiology, 2007)

Why do plants reflect in the green and have a “red edge” in the red, and should extrasolar photosynthesis be the same? We provide (1) a brief review of how photosynthesis works, (2) an overview of the diversity of photosynthetic organisms, their light harvesting systems, and environmental ranges, (3) a synthesis of photosynthetic surface spectral signatures, and (4) evolutionary rationales for photosynthetic surface reflectance spectra with regard to utilization of photon energy and the planetary light environment. We found the “near-infrared (NIR) end” of the red edge to trend from blue-shifted to reddest for (in order) snow algae, temperate algae, lichens, mosses, aquatic plants, and finally terrestrial vascular plants. The red edge is weak or sloping in lichens. Purple bacteria exhibit possibly a sloping edge in the NIR. More studies are needed on pigment–protein complexes, membrane composition, and measurements of bacteria before firm conclusions can be drawn about the role of the NIR reflectance. Pigment absorbance features are strongly correlated with features of atmospheric spectral transmittance: P680 in Photosystem II with the peak surface incident photon flux density at ∼685 nm, just before an oxygen band at 687.5 nm; the NIR end of the red edge with water absorbance bands and the oxygen A-band at 761 nm; and bacteriochlorophyll reaction center wavelengths with local maxima in atmospheric and water transmittance spectra. Given the surface incident photon flux density spectrum and resonance transfer in light harvesting, we propose some rules with regard to where photosynthetic pigments will peak in absorbance: (1) the wavelength of peak incident photon flux; (2) the longest available wavelength for core antenna or reaction center pigments; and (3) the shortest wavelengths within an atmospheric window for accessory pigments. That plants absorb less green light may not be an inefficient legacy of evolutionary history, but may actually satisfy the above criteria.

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Spectral signatures of photosynthesis. II. Coevolution with other stars and the atmosphere on extrasolar worlds (Astrobiology, 2007)

As photosynthesis on Earth produces the primary signatures of life that can be detected astronomically at the global scale, a strong focus of the search for extrasolar life will be photosynthesis, particularly photosynthesis that has evolved with a different parent star. We take previously simulated planetary atmospheric compositions for Earth-like planets around observed F2V and K2V, modeled M1V and M5V stars, and around the active M4.5V star AD Leo; our scenarios use Earth’s atmospheric composition as well as very low O2 content in case anoxygenic photosynthesis dominates. With a line-by-line radiative transfer model, we calculate the incident spectral photon flux densities at the surface of the planet and under water. We identify bands of available photosynthetically relevant radiation and find that photosynthetic pigments on planets around F2V stars may peak in absorbance in the blue, K2V in the red-orange, and M stars in the near-infrared, in bands at 0.93–1.1 μm, 1.1–1.4 μm, 1.5–1.8 μ m, and 1.8–2.5 μm.

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