BIOMEX (BIOlogy and Mars EXperiment) is an ESA/Roscosmos space exposure experiment housed within the exposure facility EXPOSE-R2 outside the Zvezda module on the International Space Station (ISS). The design of the multiuser facility supports—among others—the BIOMEX investigations into the stability and level of degradation of space-exposed biosignatures such as pigments, secondary metabolites, and cell surfaces in contact with a terrestrial and Mars analog mineral environment. In parallel, analysis on the viability of the investigated organisms has provided relevant data for evaluation of the habitability of Mars, for the limits of life, and for the likelihood of an interplanetary transfer of life (theory of lithopanspermia). In this project, lichens, archaea, bacteria, cyanobacteria, snow/permafrost algae, meristematic black fungi, and bryophytes from alpine and polar habitats were embedded, grown, and cultured on a mixture of martian and lunar regolith analogs or other terrestrial minerals. The organisms and regolith analogs and terrestrial mineral mixtures were then exposed to space and to simulated Mars-like conditions by way of the EXPOSE-R2 facility. In this special issue, we present the first set of data obtained in reference to our investigation into the habitability of Mars and limits of life. This project was initiated and implemented by the BIOMEX group, an international and interdisciplinary consortium of 30 institutes in 12 countries on 3 continents. Preflight tests for sample selection, results from ground-based simulation experiments, and the space experiments themselves are presented and include a complete overview of the scientific processes required for this space experiment and postflight analysis. The presented BIOMEX concept could be scaled up to future exposure experiments on the Moon and will serve as a pretest in low Earth orbit.

In the past decade, various astrobiological studies on different lichen species investigated the impairment of viability and photosynthetic activity by exposure to simulated or real space parameters (as vacuum, polychromatic ultraviolet (UV)-radiation and monochromatic UVC) and consistently found high post-exposure viability as well as low rates of photosynthetic impairment (de Vera et al. 2003(de Vera et al. , 2004a2004b;de la Torre et al. 2010;Onofri et al. 2012;Sánchez et al. 2012Sánchez et al. , 2014Brandt et al. 2014). To achieve a better understanding of the basic mechanisms of resistance, the present study subdued isolated and metabolically active photobionts of two astrobiologically relevant lichens to UVC 254 nm , examined its effect on photosynthetic activity by chlorophyll a fluorescence and characterized the UVC-induced damages by quantum yield reduction and measurements of non-photochemical quenching. The results indicate a strong impairment of photosynthetic activity, photoprotective mechanisms and overall photobiont vitality when being irradiated in the isolated and metabolically active state. In conclusion, the present study stresses the higher susceptibility of photobionts towards extreme environmental conditions as UVC-exposure, a stressor that does not occur on the Earth. By comparison with previous studies, the present results highlight the importance of protective mechanisms in lichens, such as morphological-anatomical traits (Meeßen et al. 2013), secondary lichen compounds ) and the symbiont's pivotal ability to pass into anhydrobiosis when desiccating.

Lichen symbioses between fungi and algae represent successful life strategies to colonize the most extreme terrestrial habitats. Consequently, space exposure and simulation experiments have demonstrated lichens' high capacity for survival, and thus, they have become models in astrobiological research with which to discern the limits and limitations of terrestrial life. In a series of ground-based irradiation experiments, the STARLIFE campaign investigated the resistance of astrobiological model organisms to galactic cosmic radiation, which is one of the lethal stressors of extraterrestrial environments. Since previous studies have identified that the alga is the more sensitive lichen symbiont, we chose the isolated photobiont Trebouxia sp. of the astrobiological model Circinaria gyrosa as a subject in the campaign. Therein, γ radiation was used to exemplify the deleterious effects of low linear energy transfer (LET) ionizing radiation at extremely high doses up to 113 kGy in the context of astrobiology. The effects were analyzed by chlorophyll a fluorescence of photosystem II (PSII), cultivation assays, live/dead staining and confocal laser scanning microscopy (CLSM), and Raman laser spectroscopy (RLS). The results demonstrate dose-dependent impairment of photosynthesis, the cessation of cell proliferation, cellular damage, a decrease in metabolic activity, and degradation of photosynthetic pigments. While previous investigations on other extraterrestrial stressors have demonstrated a high potential of resistance, results of this study reveal the limits of photobiont resistance to ionizing radiation and characterize γ radiation-induced damages. This study also supports parallel STARLIFE studies on the lichens Circinaria gyrosa and Xanthoria elegans, both of which harbor a Trebouxia sp. photobiont. Key Words: Astrobiology-Gamma rays-Extremotolerance-Ionizing radiation-Lichens-Photobiont. Astrobiology 17, 154-162.

Several investigations on lichen photobionts (PBs) after exposure to simulated or real-space parameters consistently reported high viability and recovery of photosynthetic activity. These studies focused on PBs within lichen thalli, mostly exposed in a metabolically inactive state. In contrast, a recent study exposed isolated and metabolically active PBs to the non-terrestrial stressor UVC254 nm and found strong impairment of photosynthetic activity and photo-protective mechanisms (Meeßen et al. in 2014b). Under space and Mars conditions, UVC is accompanied by other stressors as extreme desiccation and low temperatures. The present study exposed the PBs of Buellia frigida and Circinaria gyrosa, to UVC in combination with desiccation and subzero temperatures to gain better insight into the combined stressors' effect and the PBs' inherent potential of resistance. These effects were examined by chlorophyll a fluorescence which is a good indicator of photosynthetic activity (Lüttge & Büdel in 2010) and widely used to test the viability of PBs after (simulated) space exposure. The present results reveal fast recovery of photosynthetic activity after desiccation and subzero temperatures. Moreover, they demonstrate that desiccation and cold confer an additional protective effect on the investigated PBs and attenuate the PBs' reaction to another stressor – even if it is a non-terrestrial one such as UVC. Besides other protective mechanisms (anhydrobiosis, morphological–anatomical traits and secondary lichen compounds), these findings may help to explain the high resistance of lichens observed in astrobiological studies.

The lichen Buellia frigida was exposed to space and simulated Mars analog conditions in the BIOlogy and Mars EXperiment (BIOMEX) project operated outside the International Space Station (ISS) for 1.5 years. To determine the effects of the low Earth orbit (LEO) conditions on the lichen symbionts, a LIVE/DEAD staining analysis test was performed. After return from the ISS, the lichen symbionts demonstrated mortality rates of up to 100% for the algal symbiont and up to 97.8% for the fungal symbiont. In contrast, the lichen symbiont controls exhibited mortality rates of 10.3% up to 31.9% for the algal symbiont and 14.5% for the fungal symbiont. The results performed in the BIOMEX Mars simulation experiment on the ISS indicate that the potential for survival and the resistance of the lichen B. frigida to LEO conditions are very low. It is unlikely that Mars could be inhabited by this lichen, even for a limited amount of time, or even not habitable planet for the tested lichen symbionts.

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Two rover missions to Mars aim to detect biomolecules as a sign of extinct or extant life with, among other instruments, Raman spectrometers. However, there are many unknowns about the stability of Raman-detectable biomolecules in the martian environment, clouding the interpretation of the results. To quantify Raman-detectable biomolecule stability, we exposed seven biomolecules for 469 days to a simulated martian environment outside the International Space Station. Ultraviolet radiation (UVR) strongly changed the Raman spectra signals, but only minor change was observed when samples were shielded from UVR. These findings provide support for Mars mission operations searching for biosignatures in the subsurface. This experiment demonstrates the detectability of biomolecules by Raman spectroscopy in Mars regolith analogs after space exposure and lays the groundwork for a consolidated space-proven database of spectroscopy biosignatures in targeted environments.

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