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Life, Volume 8, Issue 2 (June 2018) – 16 articles

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15 pages, 694 KiB  
Review
Chemomimesis and Molecular Darwinism in Action: From Abiotic Generation of Nucleobases to Nucleosides and RNA
by Raffaele Saladino, Judit E. Šponer, Jiří Šponer, Giovanna Costanzo, Samanta Pino and Ernesto Di Mauro
Life 2018, 8(2), 24; https://doi.org/10.3390/life8020024 - 20 Jun 2018
Cited by 13 | Viewed by 5678
Abstract
Molecular Darwinian evolution is an intrinsic property of reacting pools of molecules resulting in the adaptation of the system to changing conditions. It has no a priori aim. From the point of view of the origin of life, Darwinian selection behavior, when spontaneously [...] Read more.
Molecular Darwinian evolution is an intrinsic property of reacting pools of molecules resulting in the adaptation of the system to changing conditions. It has no a priori aim. From the point of view of the origin of life, Darwinian selection behavior, when spontaneously emerging in the ensembles of molecules composing prebiotic pools, initiates subsequent evolution of increasingly complex and innovative chemical information. On the conservation side, it is a posteriori observed that numerous biological processes are based on prebiotically promptly made compounds, as proposed by the concept of Chemomimesis. Molecular Darwinian evolution and Chemomimesis are principles acting in balanced cooperation in the frame of Systems Chemistry. The one-pot synthesis of nucleosides in radical chemistry conditions is possibly a telling example of the operation of these principles. Other indications of similar cases of molecular evolution can be found among biogenic processes. Full article
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<p>Proposed mechanism of the proton irradiation induced <span class="html-italic"><span class="html-small-caps">N</span></span>-glycosidation between adenine and ribose [<a href="#B69-life-08-00024" class="html-bibr">69</a>].</p>
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<p>A ladder-like stacked supramolecular architecture provides optimum steric conditions for the oligomerization of 3′, 5′ cGMP. Left: Nucleobase stacking in the crystal structure of 3′, 5′ cGMP [<a href="#B119-life-08-00024" class="html-bibr">119</a>]. Right: Proposed structure of the trigonal bipyramidal intermediate of the chain-extension reaction from TPSS-D2/TVZP calculations [<a href="#B114-life-08-00024" class="html-bibr">114</a>]. The yellow nucleotides serve as mediators of the transphosphorylation reactions.</p>
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16 pages, 21646 KiB  
Article
Integrity of the DNA and Cellular Ultrastructure of Cryptoendolithic Fungi in Space or Mars Conditions: A 1.5-Year Study at the International Space Station
by Silvano Onofri, Laura Selbmann, Claudia Pacelli, Jean Pierre De Vera, Gerda Horneck, John E. Hallsworth and Laura Zucconi
Life 2018, 8(2), 23; https://doi.org/10.3390/life8020023 - 19 Jun 2018
Cited by 17 | Viewed by 6951
Abstract
The black fungi Cryomyces antarcticus and Cryomyces minteri are highly melanized and are resilient to cold, ultra-violet, ionizing radiation and other extreme conditions. These microorganisms were isolated from cryptoendolithic microbial communities in the McMurdo Dry Valleys (Antarctica) and studied in Low Earth Orbit [...] Read more.
The black fungi Cryomyces antarcticus and Cryomyces minteri are highly melanized and are resilient to cold, ultra-violet, ionizing radiation and other extreme conditions. These microorganisms were isolated from cryptoendolithic microbial communities in the McMurdo Dry Valleys (Antarctica) and studied in Low Earth Orbit (LEO), using the EXPOSE-E facility on the International Space Station (ISS). Previously, it was demonstrated that C. antarcticus and C. minteri survive the hostile conditions of space (vacuum, temperature fluctuations, and the full spectrum of extraterrestrial solar electromagnetic radiation), as well as Mars conditions that were simulated in space for a 1.5-year period. Here, we qualitatively and quantitatively characterize damage to DNA and cellular ultrastructure in desiccated cells of these two species, within the frame of the same experiment. The DNA and cells of C. antarcticus exhibited a higher resistance than those of C. minteri. This is presumably attributable to the thicker (melanized) cell wall of the former. Generally, DNA was readily detected (by PCR) regardless of exposure conditions or fungal species, but the C. minteri DNA had been more-extensively mutated. We discuss the implications for using DNA, when properly shielded, as a biosignature of recently extinct or extant life. Full article
(This article belongs to the Special Issue Fungi from Extreme Environments)
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<p>Cells of <span class="html-italic">Cryomyces antarcticus</span> (<b>A</b>,<b>B</b>) and <span class="html-italic">Cryomyces minteri</span> (<b>C</b>,<b>D</b>) after three months incubation on malt extract agar at 15 °C. Panels (<b>A</b>,<b>C</b>) were reproduced with permission from Selbmann et al. [<a href="#B14-life-08-00023" class="html-bibr">14</a>]. Photographs were taken using light microscopy, and scales bars = 10 μm.</p>
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<p>(<b>A</b>) Expose-E integrated in the EuTEF platform, at the Kennedy Space Center, before flight; (<b>B</b>) EXPOSE-E fitted onto the Space Shuttle; and (<b>C</b>) robotic arm in action during assembly of the EuTEF on the ISS.</p>
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<p>DNA bands obtained by single gene PCR. <span class="html-italic">C. antarcticus</span> (<b>A</b>) 700 bp; (<b>B</b>) 1600 bp; (<b>C</b>) 2000 bp and <span class="html-italic">C. minteri</span>; (<b>D</b>) 700 bp; (<b>E</b>) 1600 bp; (<b>F</b>) 2000 bp. CTR = Ground control; Space dark = space-exposed dark treatment; Space 100% = space-exposed 100% irradiation; Space 0.1% = space-exposed 0.1% irradiation; Mars dark = simulated Mars conditions, dark treatment; Mars 100% = simulated Mars conditions 100% irradiation; Mars 0.1% = simulated Mars conditions 0.1% irradiation (see <a href="#life-08-00023-t001" class="html-table">Table 1</a>); Neg = negative control and M = DNA ladder.</p>
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<p>Electropherograms showing nucleotide sequences for the ITS (60–135 position) of <span class="html-italic">C. antarcticus</span>: (<b>A</b>) Ground control; (<b>B</b>) Space 100% irradiation; (<b>C</b>) simulated Mars dark conditions; and (<b>D</b>) simulated Mars 100% irradiation.</p>
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<p>Electropherograms showing nucleotide sequences for the ITS (60–135 position) of <span class="html-italic">C. minteri</span>. (<b>A</b>) Ground control; (<b>B</b>) Space 100% irradiation; (<b>C</b>) dark Mars; and (<b>D</b>) Mars 100% irradiation.</p>
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<p>Unweighted Pair Group Method with Arithmetic Mean (UPGMA) analyses based on <span class="html-italic">C. antarcticus</span> sequences alignment; the dendrogram was rooted with the same gene sequence from the Ground control of <span class="html-italic">C. minteri</span> (<b>A</b>); UPGMA analyses based on <span class="html-italic">C. minteri</span> sequences alignment; the dendrogram rooted with Mars 100% irradiation resulted in the most mutated sequence after treatment (<b>B</b>).</p>
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<p>Genomic DNA fingerprint profiles from the RAPD assay of experimental samples (see <a href="#life-08-00023-t001" class="html-table">Table 1</a>) for: (<b>A</b>) <span class="html-italic">C. antarcticus</span> and (<b>B</b>) <span class="html-italic">C. minteri</span>. CTR = Ground control; Space dark = space-exposed dark treatment; Space 100% = space-exposed 100% irradiation; Space 0.1% = space-exposed 0.1% irradiation; Mars dark = simulated Mars conditions, dark treatment; Mars 100% = simulated Mars conditions 100% irradiation; Mars 0.1% = simulated Mars conditions 0.1% irradiation; Neg = negative control and M = DNA ladder.</p>
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<p>Number of the target gene copies according to real-time q-PCR of different treatments (see <a href="#life-08-00023-t001" class="html-table">Table 1</a>) in (<b>A</b>) <span class="html-italic">C. antarcticus</span> and (<b>B</b>) <span class="html-italic">C. minteri</span>. For different columns which have a common letter-designation above, this indicates that the values are not statistically significantly different from each other according to the <span class="html-italic">t</span> test (<span class="html-italic">p</span> ≤ 0.05). CTR = Ground control; Space dark = space-exposed dark treatment; Space 100% = space-exposed 100% irradiation; Space 0.1% = space-exposed 0.1% irradiation; Mars dark = simulated Mars conditions, dark treatment; Mars 100% = simulated Mars conditions 100% irradiation; Mars 0.1% = simulated Mars conditions 0.1% irradiation.</p>
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<p>Cells observed by TEM of the <span class="html-italic">C. antarcticus</span> Ground control (<b>A</b>,<b>B</b>), Space 100% irradiation (<b>C</b>–<b>F</b>) and <span class="html-italic">C. minteri</span> Ground control (<b>G</b>,<b>H</b>), Space 100% irradiation (<b>I</b>–<b>L</b>). Scale bars = 5 μm for (<b>B</b>,<b>G</b>,<b>H</b>,<b>J</b>–<b>L</b>) and 2 μm for (<b>A</b>,<b>C</b>–<b>F</b>,<b>I</b>).</p>
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14 pages, 2132 KiB  
Article
Big Sound and Extreme Fungi—Xerophilic, Halotolerant Aspergilli and Penicillia with Low Optimal Temperature as Invaders of Historic Pipe Organs
by Katja Sterflinger, Christian Voitl, Ksenija Lopandic, Guadalupe Piñar and Hakim Tafer
Life 2018, 8(2), 22; https://doi.org/10.3390/life8020022 - 14 Jun 2018
Cited by 16 | Viewed by 5881
Abstract
Recent investigations have shown that xerophilic fungi may pose a biodeterioration risk by threatening objects of cultural heritage including many types of materials, including wood, paint layers, organic glues or leather and even metal. Historic—and also new built—pipe organs combine all those materials. [...] Read more.
Recent investigations have shown that xerophilic fungi may pose a biodeterioration risk by threatening objects of cultural heritage including many types of materials, including wood, paint layers, organic glues or leather and even metal. Historic—and also new built—pipe organs combine all those materials. In this study, halotolerant aspergilli and penicillia with low optimal temperatures were shown to be the most frequent invaders of pipe organs. The fungi form white mycelia on the organic components of the organs with a clear preference for the bolus paint of the wooden pipes, the leather-made hinges of the stop actions and all parts fixed by organic glue. Physiological tests showed that the strains isolated from the instruments all show a halotolerant behavior, although none was halophilic. The optimum growth temperature is below 20 °C, thus the fungi are perfectly adapted to the cool and relatively dry conditions in the churches and organs respectively. The de-novo genome sequences analyses of the strains are currently ongoing and will reveal the genomic basis for the halotolerant behavior of the fungi. Full article
(This article belongs to the Special Issue Fungi from Extreme Environments)
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<p>Baroque pipe organ with decorations (Styria, Austria).</p>
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<p>Light fungal colonies on Bolus painted wood pipe.</p>
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<p>Heavy fungal deterioration of wood pipes.</p>
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<p>Fungal growth on hinges of stop actions.</p>
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<p>Fungal contamination on leather sealings (“pulpetes”) within the windchest.</p>
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<p>Fungal growth on the inner part of the organs casing.</p>
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<p>Colony morphology of the four dominant stains grown on a maximally tolerated salt concentration: (<b>A</b>) MA 6036, <span class="html-italic">Penicillium</span> sp., 20% NaCl; (<b>B</b>) MA 6037, <span class="html-italic">Aspergillus</span> sp., 25% NaCl; (<b>C</b>) MA 6039, <span class="html-italic">Aspergillus</span> sp., 15% NaCl; (<b>D</b>) MA 6040, <span class="html-italic">Penicillium rubens</span>, 25% NaCl.</p>
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14 pages, 373 KiB  
Article
Homochirality through Photon-Induced Denaturing of RNA/DNA at the Origin of Life
by Karo Michaelian
Life 2018, 8(2), 21; https://doi.org/10.3390/life8020021 - 6 Jun 2018
Cited by 10 | Viewed by 6755
Abstract
Since a racemic mixture of chiral nucleotides frustrates the enzymeless extension of RNA and DNA, the origin of homochirality must be intimately connected with the origin of life. Homochirality theories have elected to presume abiotic mechanisms for prebiotic enantiomer enrichment and post amplification, [...] Read more.
Since a racemic mixture of chiral nucleotides frustrates the enzymeless extension of RNA and DNA, the origin of homochirality must be intimately connected with the origin of life. Homochirality theories have elected to presume abiotic mechanisms for prebiotic enantiomer enrichment and post amplification, but none, so far, has been generally accepted. Here I present a novel hypothesis for the procurement of homochirality from an asymmetry in right- over left-circularly polarized photon-induced denaturing of RNA and DNA at the Archean ocean surface as temperatures descended below that of RNA and DNA melting. This asymmetry is attributed to the small excess of right-handed circularly polarized submarine light during the afternoon, when surface water temperatures were highest and thus most conducive to photon-induced denaturing, and to a negative circular dichroism band extending from 230 to 270 nm for small oligos of RNA and DNA. Because D-nucleic acids have greater affinity for L-tryptophan due to stereochemistry, and because D-RNA/DNA+L-tryptophan complexes have an increased negative circular dichroism band between 230 and 270 nm, the homochirality of tryptophan can also be explained by this hypothesis. A numerical model is presented, demonstrating the efficacy of such a mechanism in procuring homochirality of RNA or DNA from an original racemic solution in as little as 270 Archean years. Full article
(This article belongs to the Special Issue The Origin of Chirality in Life (Chiral Symmetry Breaking))
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<p>Circular dichroism of the 20 base pair oligo B-form duplex 3′-GCGGCGACTGGTGAGTACGC (M.W. 12,306 g mol<sup>−1</sup>) at neutral pH and room temperature taken from Figure 3a of [<a href="#B60-life-08-00021" class="html-bibr">60</a>] (black line with triangles). Circular dichroism of the 11 base pair hairpin B-form duplex 3′-TTTCGCGAAAT (M.W. 6839 g mol<sup>−1</sup>) (blue line with crosses, left y-scale) at neutral pH and room temperature adapted from Figure 5a of [<a href="#B61-life-08-00021" class="html-bibr">61</a>]. Circular dichroism of adenosine with tryptophan in the ratio 1:2 taken from Figure 3a of Arcaya et al. [<a href="#B62-life-08-00021" class="html-bibr">62</a>] (red line with circles: the CD spectrum for adenosine+tryptophan is plotted divided by 4 for convenience of scale. Arcaya et al. state that the CD spectrum of DNA+tryptophan is similar to the average of the CD spectra of the individual nucleosides with tryptophan [<a href="#B62-life-08-00021" class="html-bibr">62</a>]). Short oligo (25 bp) absorption spectrum taken from [<a href="#B47-life-08-00021" class="html-bibr">47</a>] normalized to a peak value of one (black dotted line, right y-scale). Note that, at wavelengths shorter than 220 nm, the absorption spectrum is not reliable due to the reduced output of the deuterium light used [<a href="#B47-life-08-00021" class="html-bibr">47</a>]. Solar Archean spectrum at Earth’s surface [<a href="#B40-life-08-00021" class="html-bibr">40</a>] normalized to a peak value of one (red dashed line, right y-scale).</p>
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<p>Circular dichroism spectra of the 20 bp oligo 3′-GCGGCGACTGGTGAGTACGC, the 11 bp oligo 3′-TTTCGCGAAAT, and adenosine+tryptophan convoluted with a short oligo (25 bp) absorption spectrum and the wavelength-dependent intensity of the light reaching Earth’s surface during the Archean (see <a href="#life-08-00021-f001" class="html-fig">Figure 1</a>). Adenosine+tryptophan is plotted at 1/4 of its real amplitude for convenience of scale. The average CD values, integrated over the region 200 to 300 nm are −0.279, −1.06 and −13.37 (220 to 300 nm) for 3′-GCGGCGACTGGTGAGTACGC, 3′-TTTCGCGAAAT and adenosine+tryptophan respectively.</p>
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<p>Chirality <span class="html-italic">C<sub>i</sub></span> as a function of the number of Archean days <span class="html-italic">i</span> for the two oligos and DNA+Tryptophan as calculated through Equation (<a href="#FD5-life-08-00021" class="html-disp-formula">5</a>). (a) The three smooth curves to the right were calculated assuming denaturation probability is simply proportional to the number of photons absorbed, but that complete denaturing occurs only in the afternoon, and a 5% excess of right-hand circularly polarized submarine light at the ocean surface during the afternoon. (b) These three curves to the left include in the calculation a stochastic photon absorption probability (±2% with respect to nominal values given by Equation (<a href="#FD2-life-08-00021" class="html-disp-formula">2</a>)) and an energy threshold for denaturation related to the complementary strand binding energy (see text).</p>
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11 pages, 894 KiB  
Technical Note
phylotaR: An Automated Pipeline for Retrieving Orthologous DNA Sequences from GenBank in R
by Dominic J. Bennett, Hannes Hettling, Daniele Silvestro, Alexander Zizka, Christine D. Bacon, Søren Faurby, Rutger A. Vos and Alexandre Antonelli
Life 2018, 8(2), 20; https://doi.org/10.3390/life8020020 - 5 Jun 2018
Cited by 23 | Viewed by 8805
Abstract
The exceptional increase in molecular DNA sequence data in open repositories is mirrored by an ever-growing interest among evolutionary biologists to harvest and use those data for phylogenetic inference. Many quality issues, however, are known and the sheer amount and complexity of data [...] Read more.
The exceptional increase in molecular DNA sequence data in open repositories is mirrored by an ever-growing interest among evolutionary biologists to harvest and use those data for phylogenetic inference. Many quality issues, however, are known and the sheer amount and complexity of data available can pose considerable barriers to their usefulness. A key issue in this domain is the high frequency of sequence mislabeling encountered when searching for suitable sequences for phylogenetic analysis. These issues include, among others, the incorrect identification of sequenced species, non-standardized and ambiguous sequence annotation, and the inadvertent addition of paralogous sequences by users. Taken together, these issues likely add considerable noise, error or bias to phylogenetic inference, a risk that is likely to increase with the size of phylogenies or the molecular datasets used to generate them. Here we present a software package, phylotaR that bypasses the above issues by using instead an alignment search tool to identify orthologous sequences. Our package builds on the framework of its predecessor, PhyLoTa, by providing a modular pipeline for identifying overlapping sequence clusters using up-to-date GenBank data and providing new features, improvements and tools. We demonstrate and test our pipeline’s effectiveness by presenting trees generated from phylotaR clusters for two large taxonomic clades: Palms and primates. Given the versatility of this package, we hope that it will become a standard tool for any research aiming to use GenBank data for phylogenetic analysis. Full article
(This article belongs to the Special Issue Open Science Phyloinformatics: Resources, Methods, and Analyses)
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<p>The phylotaR pipeline identifies all sequences in GenBank associated with a user-specified taxonomic identity (<b>a</b>). The pipeline then performs all-vs.-all BLAST across all the sequences to identify orthologous clusters (<b>b</b>). These searches are constrained to run within taxonomic groups up to a user-determined limit (default 50,000 sequences and 100,000 nodes). To generate higher taxonomic level clusters, an additional BLAST search is performed of the most connected sequences within clusters (i.e., the seed sequences) from the lower-level clusters. The clusters of overlapping seed sequences are then merged into larger clusters (<b>c</b>). All clusters, merged and non-merged, are then reported for inspection by the user. For more details on the pipeline, see <a href="#app2-life-08-00020" class="html-app">Appendix A</a>.</p>
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<p>Initiating the phylotaR pipeline in R for primates (TaxID: 9443).</p>
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<p>Presence/absence of tribes and genera for palms (<b>a</b>) and primates (<b>b</b>), respectively, across the top ten best clusters. X-axis numbers are unique cluster Ids. For more details on each of these clusters, see <a href="#app1-life-08-00020" class="html-app">Tables S4a and S4b</a>.</p>
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<p>Tribe- and genus-level trees for palms (<b>a</b>) and primates (<b>b</b>). Roots were determined manually by rooting with <span class="html-italic">Strepsirrhini</span> and <span class="html-italic">Calamoideae</span> for primates and palms respectively. Branch lengths have been removed. Support calculated from 100 rapid bootstraps: *** &gt;0.95, ** &gt;0.75 and * &gt;0.50. Complete tree construction methods are in <a href="#app3-life-08-00020" class="html-app">Appendix B</a>. For tree comparisons with published trees for palms [<a href="#B37-life-08-00020" class="html-bibr">37</a>] and primates [<a href="#B38-life-08-00020" class="html-bibr">38</a>], see <a href="#app1-life-08-00020" class="html-app">Figures S4 and S5</a>.</p>
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15 pages, 2410 KiB  
Article
Sun Exposure Shapes Functional Grouping of Fungi in Cryptoendolithic Antarctic Communities
by Claudia Coleine, Laura Zucconi, Silvano Onofri, Nuttapon Pombubpa, Jason E. Stajich and Laura Selbmann
Life 2018, 8(2), 19; https://doi.org/10.3390/life8020019 - 2 Jun 2018
Cited by 36 | Viewed by 6496
Abstract
Antarctic cryptoendolithic microbial communities dominate ice-free areas of continental Antarctica, among the harshest environments on Earth. The endolithic lifestyle is a remarkable adaptation to the exceptional environmental extremes of this area, which is considered the closest terrestrial example to conditions on Mars. Recent [...] Read more.
Antarctic cryptoendolithic microbial communities dominate ice-free areas of continental Antarctica, among the harshest environments on Earth. The endolithic lifestyle is a remarkable adaptation to the exceptional environmental extremes of this area, which is considered the closest terrestrial example to conditions on Mars. Recent efforts have attempted to elucidate composition of these extremely adapted communities, but the functionality of these microbes have remained unexplored. We have tested for interactions between measured environmental characteristics, fungal community membership, and inferred functional classification of the fungi present and found altitude and sun exposure were primary factors. Sandstone rocks were collected in Victoria Land, Antarctica along an altitudinal gradient from 834 to 3100 m a.s.l.; differently sun-exposed rocks were selected to test the influence of this parameter on endolithic settlement. Metabarcoding targeting the fungal internal transcribed spacer region 1 (ITS1) was used to catalogue the species found in these communities. Functional profile of guilds found in the samples was associated to species using FUNGuild and variation in functional groups compared across sunlight exposure and altitude. Results revealed clear dominance of lichenized and stress-tolerant fungi in endolithic communities. The main variations in composition and abundance of functional groups among sites correlated to sun exposure, but not to altitude. Full article
(This article belongs to the Special Issue Fungi from Extreme Environments)
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<p>Localities visited in the McMurdo Dry Valleys (Southern Victoria Land), showing different sun exposition: (<b>A</b>,<b>B</b>) Battleship Promontory North and South, respectively; (<b>C</b>,<b>D</b>) University Valley North and South, respectively; (<b>E</b>,<b>F</b>) Siegfried Peak North and South, respectively; (<b>G</b>,<b>H</b>) Finger Mt. North and South, respectively.</p>
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<p>Spearman’s correlation coefficients between fungal richness of each functional group along an altitudinal gradient. <span class="html-italic">p &gt;</span> 0.05 in LIC, SAP and SAP + PP panels; <span class="html-italic">p</span> &lt; 0.05 in RIF + BY panel.</p>
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<p>Spearman’s correlation coefficients between fungal biodiversity (Shannon’s index) of each functional group along an altitudinal gradient. <span class="html-italic">p &gt;</span> 0.05 in all four panels.</p>
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<p>Non-metric multidimensional scaling (NMDS) ordination plots for each functional fungal group of Antarctic endolithic communities differently sun exposed (blue lines: south sun exposition; red lines: north sun exposition), based on square-root transformed abundance data.</p>
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<p>Venn diagram of the four functional groups of fungi showing the distribution of OTUs between north and south exposition. The abbreviations represent functional groups. Both the percentages of OTUs that were shared and found exclusively in each sun exposure are indicated.</p>
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31 pages, 1809 KiB  
Review
Data-Driven Astrochemistry: One Step Further within the Origin of Life Puzzle
by Alexander Ruf, Louis L. S. D’Hendecourt and Philippe Schmitt-Kopplin
Life 2018, 8(2), 18; https://doi.org/10.3390/life8020018 - 1 Jun 2018
Cited by 29 | Viewed by 10539
Abstract
Astrochemistry, meteoritics and chemical analytics represent a manifold scientific field, including various disciplines. In this review, clarifications on astrochemistry, comet chemistry, laboratory astrophysics and meteoritic research with respect to organic and metalorganic chemistry will be given. The seemingly large number of observed astrochemical [...] Read more.
Astrochemistry, meteoritics and chemical analytics represent a manifold scientific field, including various disciplines. In this review, clarifications on astrochemistry, comet chemistry, laboratory astrophysics and meteoritic research with respect to organic and metalorganic chemistry will be given. The seemingly large number of observed astrochemical molecules necessarily requires explanations on molecular complexity and chemical evolution, which will be discussed. Special emphasis should be placed on data-driven analytical methods including ultrahigh-resolving instruments and their interplay with quantum chemical computations. These methods enable remarkable insights into the complex chemical spaces that exist in meteorites and maximize the level of information on the huge astrochemical molecular diversity. In addition, they allow one to study even yet undescribed chemistry as the one involving organomagnesium compounds in meteorites. Both targeted and non-targeted analytical strategies will be explained and may touch upon epistemological problems. In addition, implications of (metal)organic matter toward prebiotic chemistry leading to the emergence of life will be discussed. The precise description of astrochemical organic and metalorganic matter as seeds for life and their interactions within various astrophysical environments may appear essential to further study questions regarding the emergence of life on a most fundamental level that is within the molecular world and its self-organization properties. Full article
(This article belongs to the Special Issue Meteorites and the Origin of Life)
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<p>Sketching chemical evolution in terms of molecular diversity and molecular complexity. Molecular transformation within time and space is illustrated. Simple molecules within interstellar and circumstellar media evolve to highly-oriented, organized, complex macromolecules on planetary systems, enabling the potential of living systems.</p>
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<p>Classification of meteorites. This figure is adapted from the classification scheme, as shown by Weisberg, McCoy and Krot [<a href="#B52-life-08-00018" class="html-bibr">52</a>].</p>
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<p>Model of the molecular structure of Murchison insoluble organic matter. The figure is adapted with permission from John Wiley &amp; Sons, Inc., Hoboken, New Jersey, United States [<a href="#B76-life-08-00018" class="html-bibr">76</a>].</p>
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<p>Van Krevelen diagram of Murchison soluble organic matter. O/C versus H/C is plotted for negative ionization ESI-FT-ICR-MS methanolic soluble organic matter of Murchison. The bubble size is normalized to mass spectrometric intensity. Chemical subspaces: CHO (blue), CHNO (orange), CHOS (green), CHNOS (red) with its respective partitions. Approximately 15,000 molecular formulae are shown [<a href="#B20-life-08-00018" class="html-bibr">20</a>].</p>
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<p>Mass difference network of Murchison soluble organic matter, as analyzed from negative ionization ESI-FT-ICR-MS data. The network was visualized via Gephi software [<a href="#B141-life-08-00018" class="html-bibr">141</a>], using the Force Atlas2 layout algorithm. Chemical subspaces: CHO (blue), CHNO (orange), CHOS (green), CHNOS (red) with its respective partitions. Approximately 15,000 molecular formulae are shown [<a href="#B20-life-08-00018" class="html-bibr">20</a>].</p>
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<p>Model of the molecular structure of Murchison soluble organic matter. The figure is adapted with permission from John Wiley &amp; Sons, Inc., Hoboken, New Jersey, United States [<a href="#B150-life-08-00018" class="html-bibr">150</a>].</p>
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<p>Curiosity-driven research. This figure is adapted with permission from ©The Nobel Foundation, Nobel lecture of Laureate Theodor W. Hänsch, Stockholm, 8 December 2005 [<a href="#B156-life-08-00018" class="html-bibr">156</a>].</p>
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<p>Role of nitrogen and sulfur chemistry. (<b>A</b>) The thermally- and shock-stressed Chelyabinsk showed high numbers of nitrogen atoms within CHNO molecular formulas [<a href="#B133-life-08-00018" class="html-bibr">133</a>]; (<b>B</b>) The extremely thermally-altered Sutter’s mill reflects a loss in organic diversity, but an increase in the polysulfur domain, as compared to Murchison [<a href="#B21-life-08-00018" class="html-bibr">21</a>,<a href="#B166-life-08-00018" class="html-bibr">166</a>]. The figure is adapted with permission from The American Association for the Advancement of Science, Washington, D.C., United States [<a href="#B133-life-08-00018" class="html-bibr">133</a>,<a href="#B166-life-08-00018" class="html-bibr">166</a>].</p>
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<p>Metal distributions for meteorites, Earth and a biotic system. Data are adapted from Lodders for CI chondrites [<a href="#B50-life-08-00018" class="html-bibr">50</a>], McDonough for Earth [<a href="#B196-life-08-00018" class="html-bibr">196</a>] and Williams for cell cytoplasm [<a href="#B197-life-08-00018" class="html-bibr">197</a>].</p>
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14 pages, 2928 KiB  
Article
Testing the Domino Theory of Gene Loss in Buchnera aphidicola: The Relevance of Epistatic Interactions
by David J. Martínez-Cano, Gil Bor, Andrés Moya and Luis Delaye
Life 2018, 8(2), 17; https://doi.org/10.3390/life8020017 - 29 May 2018
Cited by 3 | Viewed by 5068
Abstract
The domino theory of gene loss states that when some particular gene loses its function and cripples a cellular function, selection will relax in all functionally related genes, which may allow for the non-functionalization and loss of these genes. Here we study the [...] Read more.
The domino theory of gene loss states that when some particular gene loses its function and cripples a cellular function, selection will relax in all functionally related genes, which may allow for the non-functionalization and loss of these genes. Here we study the role of epistasis in determining the pattern of gene losses in a set of genes participating in cell envelope biogenesis in the endosymbiotic bacteria Buchnera aphidicola. We provide statistical evidence indicating pairs of genes in B. aphidicola showing correlated gene loss tend to have orthologs in Escherichia coli known to have alleviating epistasis. In contrast, pairs of genes in B. aphidicola not showing correlated gene loss tend to have orthologs in E. coli known to have aggravating epistasis. These results suggest that during the process of genome reduction in B. aphidicola by gene loss, positive or alleviating epistasis facilitates correlated gene losses while negative or aggravating epistasis impairs correlated gene losses. We interpret this as evidence that the reduced proteome of B. aphidicola contains less pathway redundancy and more compensatory interactions, mimicking the situation of E. coli when grown under environmental constrains. Full article
(This article belongs to the Special Issue Evolution of Mutualistic Symbiosis)
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Graphical abstract

Graphical abstract
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<p>Functional classification of 102 genes from <span class="html-italic">E</span>. <span class="html-italic">coli</span> with orthologs in <span class="html-italic">Buchnera</span>. Each cell denotes the number of genes classified in each COG category. We also show the localization of the corresponding protein product in four cellular compartments. The classification of these genes follows that of Babu et al. (2011).</p>
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<p>Distribution of number of gene losses per gene family. For each one of the 102 gene families studied here, we counted the number of gene losses among the 18 <span class="html-italic">Buchnera</span> genomes.</p>
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<p>Distribution of gene losses among <span class="html-italic">Buchnera</span> genomes. The number of gene losses in the 102 gene families studied here is unevenly distributed among <span class="html-italic">Buchnera</span> genomes. The size of the circles corresponds to the number of gene losses.</p>
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<p>Gene families showing correlated gene losses along <span class="html-italic">Buchnera</span> phylogeny. Doted lines: Correlated gene losses showing a <span class="html-italic">p</span>-value &lt; 0.05. Solid lines: Correlated gene losses discovered by setting a FDR &lt; 0.1. Green lines: Positive epistatic interactions in minimal medium. Red lines: Negative epistatic interactions in minimal medium. Epistatic interactions are according to Babu et al. (2011). For clarity, epistatic interactions are shown only for those correlated gene losses showing an FDR &lt; 0.1. An FDR &lt; 0.1 captures all interactions with a <span class="html-italic">p</span>-value &lt; 0.0015.</p>
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<p>Localization of genes showing correlated gene loss (FDR &lt; 0.1) along <span class="html-italic">Buchnera</span> genomes. The size of the genomes is shown with horizontal lines and the relative localization of the genes showing correlated gene lose with colored squares. The vertical lines in each one of the genes indicate the direction of transcription. For clarity, the genome of <span class="html-italic">E</span>. <span class="html-italic">coli</span> is not shown. None of the genes are coded contiguously. <span class="html-italic">surA</span> and <span class="html-italic">ispH</span> look contiguous because of the scale, but are separated by more than 7000 bp.</p>
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<p>Comparison of correlated gene loses (CGL) identified in <span class="html-italic">Buchnera</span> with epistatic interactions in <span class="html-italic">E</span>. <span class="html-italic">coli</span> detected by Babu et al. (2011). (<b>a</b>) Dark blue triangle: All <span class="html-italic">E</span>. <span class="html-italic">coli</span> gen pairs are accommodated in an n x n matrix (only “half” the matrix is needed, below its main diagonal). Light blue triangle: Pairs of genes studied by Babu et al. (2011). Gold triangle: Pairs of <span class="html-italic">E</span>. <span class="html-italic">coli</span> genes with orthologs in <span class="html-italic">Buchnera</span> studied here. Gray triangle: Pairs of genes showing correlated gene lose in <span class="html-italic">Buchnera</span>. Dashed triangle: Pairs of genes showing epistatic interactions by Babu et al. (2011). Among the set of genes showing epistatic interactions in <span class="html-italic">E</span>. <span class="html-italic">coli</span>, some have orthologs in <span class="html-italic">Buchnera</span> showing CGL and some of them do not. (<b>b</b>) Frequency of gene families showing positive and negative epistatic interactions and showing (or not) CGL. The frequency is shown for an FDR &lt; 0.45 and for a FDR &lt; 0.1. The frequency is also shown for minimal and rich media. An FDR &lt; 0.45 comprises all CGL showing a <span class="html-italic">p</span>-value &lt; 0.05. The size of the triangles is not to scale.</p>
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<p>Ratio of positive versus negative epistatic interactions for pairs of essential (E) genes and pairs of non-essential (Non-E) genes in rich and minimal medium. The epistasis data is from <span class="html-italic">E</span>. <span class="html-italic">coli</span> genes having clear orthologs in <span class="html-italic">Buchnera</span> genomes.</p>
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15 pages, 2082 KiB  
Article
Molecular Evolution in a Peptide-Vesicle System
by Christian Mayer, Ulrich Schreiber, María J. Dávila, Oliver J. Schmitz, Amela Bronja, Martin Meyer, Julia Klein and Sven W. Meckelmann
Life 2018, 8(2), 16; https://doi.org/10.3390/life8020016 - 24 May 2018
Cited by 31 | Viewed by 14259
Abstract
Based on a new model of a possible origin of life, we propose an efficient and stable system undergoing structural reproduction, self-optimization, and molecular evolution. This system is being formed under realistic conditions by the interaction of two cyclic processes, one of which [...] Read more.
Based on a new model of a possible origin of life, we propose an efficient and stable system undergoing structural reproduction, self-optimization, and molecular evolution. This system is being formed under realistic conditions by the interaction of two cyclic processes, one of which offers vesicles as the structural environment, with the other supplying peptides from a variety of amino acids as versatile building blocks. We demonstrate that structures growing in a combination of both cycles have the potential to support their own existence, to undergo chemical and structural evolution, and to develop unpredicted functional properties. The key mechanism is the mutual stabilization of the peptides by the vesicles and of the vesicles by the peptides together with a constant production and selection of both. The development of the proposed system over time would not only represent one of the principles of life, but could also be a model for the formation of self-evolving structures ultimately leading to the first living cell. The experiment yields clear evidence for a vesicle-induced accumulation of membrane-interacting peptide which could be identified by liquid chromatography combined with high-resolution mass spectroscopy. We found that the selected peptide has an immediate effect on the vesicles, leading to (i) reduced vesicle size, (ii) increased vesicle membrane permeability, and (iii) improved thermal vesicle stability. Full article
(This article belongs to the Section Hypotheses in the Life Sciences)
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Figure 1
<p>Mechanism of peptide selection and accumulation in presence of vesicles. <b>Left</b>: Peptide chains formed by hydrophilic amino acids (blue circles) will undergo little interaction with vesicles and remain in the aqueous phase where they undergo hydrolysis. <b>Right</b>: Peptide chains formed by hydrophobic amino acids (red circles) will eventually be eluted by scCO<sub>2</sub>. <b>Center</b>: Amphiphilic peptides will accumulate in the bilayer membrane and remain partially protected against hydrolysis and elution [<a href="#B16-life-08-00016" class="html-bibr">16</a>]. The thickness of the vesicle membrane is exaggerated in order to visualize the internal structure. (Figure reproduced from ref. [<a href="#B16-life-08-00016" class="html-bibr">16</a>]).</p>
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<p>Optical micrograph of the vesicle dispersion in a sample taken during the described evolution experiment. The picture was produced under phase-contrast illumination in order to increase the visibility of the vesicles. The small fraction of vesicles larger than 2 µm can be resolved, and their membrane structure becomes apparent (arrows). Other particles in the image are primarily crystallized components. The picture was taken on a sample with a vesicle concentration of approximately 1 vol %, at neutral pH and at room temperature.</p>
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<p>Stejskal-Tanner-plot of PFG-NMR data obtained on water molecules in a sample of the aqueous phase during an evolution experiment. The steep initial part of the echo decay corresponds to free water, and the following shallow part to encapsulated water inside the vesicles. The values for ∆ (25, 50, and 100 ms) refer to the spacing between the gradient pulses. The measurement was obtained on a sample with a vesicle concentration of approximately 0.5 vol %, at neutral pH and at room temperature.</p>
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<p>Principal component analysis of samples from an evolution experiment taken after 0 and 160 h. The red box labels the entity of molecules which are absent at t = 0 h and only form in the presence of vesicles over time.</p>
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<p>Extracted ion chromatogram of the selected mass peak representing the octapeptide composition. H-Lys-Ser-Pro-Phe-Pro-Phe-Ala-Ala-OH (<span class="html-italic">m/z</span> = 863.4548 g/mol for the non-protonated species). The three signals (two strong and one weak) presumably correspond to different amino acid sequences with the same overall composition.</p>
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<p>Average diameter of the vesicles before (<b>left</b>) and after the addition of the peptide H-Lys-Ser-Pro-Phe-Pro-Phe-Ala-Ala-OH (<b>right</b> of dotted line). The error bars mark the experimental variability.</p>
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<p>Stejskal-Tanner plots determined on water for neat vesicles (top row: <b>a</b>–<b>c</b>) and after the addition of the peptide H-Lys-Ser-Pro-Phe-Pro-Phe-Ala-Ala-OH (bottom row: <b>d</b>–<b>f</b>). As determined from the inversion of the sequence for ∆ = 25, 50, and 100 ms, the peptide causes a temporary increase of the vesicle membrane permeability for water (<b>d</b>). Within several hours, the sequence slowly returns to the original one, indicating the loss of the original permeability over time (<b>e</b>,<b>f</b>).</p>
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<p>Stejskal-Tanner plots determined on water for neat vesicles (top row, <b>a</b>–<b>c</b>) and after the addition of the peptide H-Lys-Ser-Pro-Phe-Pro-Phe-Ala-Ala-OH (bottom row: <b>d</b>–<b>f</b>). Both series correspond to storage at 50 °C for 0, 19, and 100 h. As determined from the shift of the final plot level, there is a significant decomposition of the neat vesicles (top row: <b>a</b>–<b>c</b>). No such decomposition is observed on vesicles containing the peptide, revealing its stabilizing effect.</p>
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<p>Possible targets of a selection process of peptide-vesicle systems. Integration and protection of peptide (parasitic), additional thermal stabilization of the vesicle structure (symbiotic), and introduction of a peptide-induced function (functional). The thickness of the vesicle membrane is exaggerated in order to visualize the internal structure.</p>
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<p>Schematic diagram of a possible structural evolution of vesicles by repeated optimization steps in tectonic fault zones. The thickness of the vesicle membrane is exaggerated in order to visualize the internal structure.</p>
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14 pages, 1065 KiB  
Review
Fungal Diversity in Lichens: From Extremotolerance to Interactions with Algae
by Lucia Muggia and Martin Grube
Life 2018, 8(2), 15; https://doi.org/10.3390/life8020015 - 22 May 2018
Cited by 73 | Viewed by 13147
Abstract
Lichen symbioses develop long-living thallus structures even in the harshest environments on Earth. These structures are also habitats for many other microscopic organisms, including other fungi, which vary in their specificity and interaction with the whole symbiotic system. This contribution reviews the recent [...] Read more.
Lichen symbioses develop long-living thallus structures even in the harshest environments on Earth. These structures are also habitats for many other microscopic organisms, including other fungi, which vary in their specificity and interaction with the whole symbiotic system. This contribution reviews the recent progress regarding the understanding of the lichen-inhabiting fungi that are achieved by multiphasic approaches (culturing, microscopy, and sequencing). The lichen mycobiome comprises a more or less specific pool of species that can develop symptoms on their hosts, a generalist environmental pool, and a pool of transient species. Typically, the fungal classes Dothideomycetes, Eurotiomycetes, Leotiomycetes, Sordariomycetes, and Tremellomycetes predominate the associated fungal communities. While symptomatic lichenicolous fungi belong to lichen-forming lineages, many of the other fungi that are found have close relatives that are known from different ecological niches, including both plant and animal pathogens, and rock colonizers. A significant fraction of yet unnamed melanized (‘black’) fungi belong to the classes Chaethothyriomycetes and Dothideomycetes. These lineages tolerate the stressful conditions and harsh environments that affect their hosts, and therefore are interpreted as extremotolerant fungi. Some of these taxa can also form lichen-like associations with the algae of the lichen system when they are enforced to symbiosis by co-culturing assays. Full article
(This article belongs to the Special Issue Fungi from Extreme Environments)
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Figure 1
<p>Habit of lichenicolous fungi on their lichen host. (<b>a</b>) <span class="html-italic">Tremella</span> sp. on <span class="html-italic">Cladonia furcata</span>; (<b>b</b>) A, <span class="html-italic">Rhagadostoma lichenicola</span> on <span class="html-italic">Solorina crocea</span>; (<b>c</b>) <span class="html-italic">Sagediopsis fissurisedens</span> on <span class="html-italic">Aspilidea myrinii</span>; (<b>d</b>) <span class="html-italic">Sclerococcum sphaerale</span> on <span class="html-italic">Pertusaria corallina</span>; (<b>e</b>) <span class="html-italic">Endococcus perpusillus</span> on <span class="html-italic">Schaereria fuscocinerea</span>; (<b>f</b>) <span class="html-italic">Rosellinula haplospora</span> on <span class="html-italic">Aspicilia caesiocinerea</span>; (<b>g</b>) <span class="html-italic">Lichenodiplis lecanorae</span> on <span class="html-italic">Tephromela atra</span> (stars labelling the pycnidia of the lichen mycobiont); (<b>h</b>) <span class="html-italic">Minutoexcipula tuerkii</span> on <span class="html-italic">Pertusaria glomerata</span>; and (<b>i</b>) detail of sporodochium and conidia (arrow) of <span class="html-italic">Minutoexcipula tuerkii</span> on <span class="html-italic">Pertusaria glomerata</span>. Arrows point to the perithecia (<b>b</b>,<b>c</b>,<b>e</b>,<b>f</b>,) and sporodochia (<b>d</b>,<b>g</b>,<b>h</b>) of the lichenicolous fungi. Scale bars: (<b>a</b>,<b>b</b>) = 2 mm, (<b>c</b>) = 1 mm, (<b>d</b>–<b>h</b>) = 0.5 mm, and (<b>i</b>) = 20 μm.</p>
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<p>Schematic phylogenetic representation of major lineages in which lichen-associated fungi are found. The phylogeny was graphically reconstructed merging information from most recent phylogenetic studies [<a href="#B64-life-08-00015" class="html-bibr">64</a>,<a href="#B66-life-08-00015" class="html-bibr">66</a>,<a href="#B67-life-08-00015" class="html-bibr">67</a>,<a href="#B68-life-08-00015" class="html-bibr">68</a>,<a href="#B69-life-08-00015" class="html-bibr">69</a>].</p>
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<p>Habit of axenically isolated lichen-associated fungi. The sample ID, its phylogenetic placement (class and/or order or lineage, sensu Muggia et al. [<a href="#B15-life-08-00015" class="html-bibr">15</a>,<a href="#B65-life-08-00015" class="html-bibr">65</a>]) and the acronym of the medium on which it grows are reported. (<b>a</b>) A1085, Leotiomycetes, DG18; (<b>b</b>) A1148, Eurotiomycetes, Chaetothyriales, clade VI, SAB; (<b>c</b>) A1073, Dothideomycetes, Myriangiales, DG18; (<b>d</b>) A1153, Eurotiomycetes, <span class="html-italic">Sclerococcum</span>-clade, LBM; (<b>e</b>) A1109, Eurotiomycetes, Chaetothyriales, clade VI, MY; and (<b>f</b>) A1074, Dothideomycetes, Pleosporales, DG18. The different colours of the mycelia in B and C are derived from a variation in melanization and belong to the same fungus. Growth medium acronyms: DG18, Dichloran/Glycerol agar [<a href="#B80-life-08-00015" class="html-bibr">80</a>]; LBM, Lilly &amp; Barnett medium [<a href="#B81-life-08-00015" class="html-bibr">81</a>]; MY, Malt Yeast-extract [<a href="#B81-life-08-00015" class="html-bibr">81</a>]; SAB, Sabouraud [<a href="#B82-life-08-00015" class="html-bibr">82</a>]. Scale bars: (<b>a</b>–<b>d</b>,<b>f</b>) = 4 mm, and (<b>e</b>) = 2 mm.</p>
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21 pages, 3709 KiB  
Review
Insights into Abiotically-Generated Amino Acid Enantiomeric Excesses Found in Meteorites
by Aaron S. Burton and Eve L. Berger
Life 2018, 8(2), 14; https://doi.org/10.3390/life8020014 - 12 May 2018
Cited by 43 | Viewed by 10793
Abstract
Biology exhibits homochirality, in that only one of two possible molecular configurations (called enantiomers) is used in both proteins and nucleic acids. The origin of this phenomenon is currently unknown, as nearly all known abiotic mechanisms for generating these compounds result in equal [...] Read more.
Biology exhibits homochirality, in that only one of two possible molecular configurations (called enantiomers) is used in both proteins and nucleic acids. The origin of this phenomenon is currently unknown, as nearly all known abiotic mechanisms for generating these compounds result in equal (racemic) mixtures of both enantiomers. However, analyses of primitive meteorites have revealed that a number of amino acids of extraterrestrial origin are present in enantiomeric excess, suggesting that there was an abiotic route to synthesize amino acids in a non-racemic manner. Here we review the amino acid contents of a range of meteorites, describe mechanisms for amino acid formation and their potential to produce amino acid enantiomeric excesses, and identify processes that could have amplified enantiomeric excesses. Full article
(This article belongs to the Special Issue Meteorites and the Origin of Life)
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Figure 1

Figure 1
<p>Structural drawings of amino acids and sugars. (<b>a</b>) A generic α-amino acid, where the amino group is connected to the carbon immediately adjacent to the carboxylic acid. (<b>b</b>) Stereorepresentations of the two enantiomers of a chiral α-amino acid, denoted S and R. (<b>c</b>) Fischer projections denoting chirality of a chiral α-amino acid and sugar (chiral carbons marked with asterisks), where L denotes left and D denotes right; this naming convention only reflects the first chiral center.</p>
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<p>Total amino acid abundances (bars, primary axis) and percentage of five-carbon amino acids where the amino group is attached to the carbon adjacent to the carboxylic acid (α-amino acids; open circles, secondary axis) among the carbonaceous chondrite groups [<a href="#B15-life-08-00014" class="html-bibr">15</a>,<a href="#B33-life-08-00014" class="html-bibr">33</a>,<a href="#B34-life-08-00014" class="html-bibr">34</a>,<a href="#B35-life-08-00014" class="html-bibr">35</a>,<a href="#B36-life-08-00014" class="html-bibr">36</a>,<a href="#B37-life-08-00014" class="html-bibr">37</a>,<a href="#B38-life-08-00014" class="html-bibr">38</a>].</p>
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<p>Amino acids that have been found in enantiomeric excesses &gt;2% in multiple meteorites. Numbers in parentheses denote the highest values reported [<a href="#B37-life-08-00014" class="html-bibr">37</a>,<a href="#B50-life-08-00014" class="html-bibr">50</a>,<a href="#B57-life-08-00014" class="html-bibr">57</a>,<a href="#B62-life-08-00014" class="html-bibr">62</a>].</p>
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<p>Isovaline enantiomeric excesses of 0 to 20.5% have been reported [<a href="#B14-life-08-00014" class="html-bibr">14</a>,<a href="#B37-life-08-00014" class="html-bibr">37</a>,<a href="#B54-life-08-00014" class="html-bibr">54</a>,<a href="#B61-life-08-00014" class="html-bibr">61</a>,<a href="#B62-life-08-00014" class="html-bibr">62</a>]. Antarctic meteorite abbreviations are Allan Hills (ALH), Scott Glaciers (SCO), Miller Range (MIL), Pecora Escarpment (PCA), Grosvenor Mountains (GRO), Patuxent Range (PAT), Lewis Cliff (LEW) and Queen Alexandria Range (QUE). The value for Murchison reflects the highest reported, but isovaline has been detected in 0 to 18% enantiomeric excess in Murchison.</p>
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<p>Illustration of α-amino acid synthesis routes that could have taken place in meteorite parent bodies. (<b>a</b>) The Strecker-cyanohydrin pathway relies on the reaction of aldehydes or ketones with ammonia, cyanide and water to produce amino acids. (<b>b</b>) Reductive amination reactions involve the reaction of α-keto acids with ammonia and a reductant in the meteorite parent body to produce amino acids.</p>
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<p>The absolute configuration (R or S) of a chiral molecule is based on the arrangement of atoms directly attached to the stereocenter. For the <span class="html-small-caps">l</span>-enantiomers of the amino acids listed above, all but cysteine have an absolute configuration of S. The optical activity of chiral molecules describe the direction in which each enantiomer rotates plane polarized light; clockwise is dextrorotatory (+), counterclockwise direction is levorotatory (−). The sign of the optical rotation of an enantiomer is not directly related to its absolute configuration, and can vary based on the solution in which it is measured. In addition to being solution-dependent, the optical rotation varies with the molarity of each solution. For many of the amino acids, the entire range of specific rotation values fall in the positive or negative realm; exceptions to this include <span class="html-small-caps">l</span>-Glutamic Acid and <span class="html-small-caps">l</span>-Tryptophan, which have specific rotation values that vary between negative and positive numbers in specific solvents. <span class="html-small-caps">l</span>-cysteine, <span class="html-small-caps">l</span>-isoleucine, and <span class="html-small-caps">l</span>-<span class="html-italic">allo</span>-isoleucine are dextrorotatory in glacial acetic acid. References a-k correspond to [<a href="#B82-life-08-00014" class="html-bibr">82</a>,<a href="#B83-life-08-00014" class="html-bibr">83</a>,<a href="#B84-life-08-00014" class="html-bibr">84</a>,<a href="#B85-life-08-00014" class="html-bibr">85</a>,<a href="#B86-life-08-00014" class="html-bibr">86</a>,<a href="#B87-life-08-00014" class="html-bibr">87</a>,<a href="#B88-life-08-00014" class="html-bibr">88</a>,<a href="#B89-life-08-00014" class="html-bibr">89</a>,<a href="#B90-life-08-00014" class="html-bibr">90</a>,<a href="#B91-life-08-00014" class="html-bibr">91</a>,<a href="#B92-life-08-00014" class="html-bibr">92</a>,<a href="#B93-life-08-00014" class="html-bibr">93</a>].</p>
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<p>Schematic of crystallization-based enantioenrichment mechanisms. (<b>a</b>) For amino acids that preferentially form racemic crystals, in a saturated solution with one enantiomer present in excess, the solid phase would be racemic and the solution phase would be enriched to a greater degree than the overall enantiomeric excess (counting both the solution and solid phase). The enantioenriched solution phase can be physically separated from the racemic solid during aqueous alteration. (<b>b</b>) For amino acids that preferentially form enantiopure crystals, in a saturated solution where one enantiomer is present in excess, that enantiomer will tend to form more crystals that are larger, while the less abundant enantiomer will form fewer crystals that are smaller. In this case the solution phase will be racemic, so racemization would affect both enantiomers equally. However, producing more of the more abundant enantiomer will cause more of it to precipitate, increase the enantiomeric excess.</p>
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15 pages, 3249 KiB  
Article
Caveats to Exogenous Organic Delivery from Ablation, Dilution, and Thermal Degradation
by Chris Mehta, Anthony Perez, Glenn Thompson and Matthew A. Pasek
Life 2018, 8(2), 13; https://doi.org/10.3390/life8020013 - 12 May 2018
Cited by 16 | Viewed by 5441
Abstract
A hypothesis in prebiotic chemistry argues that organics were delivered to the early Earth in abundance by meteoritic sources. This study tests that hypothesis by measuring how the transfer of organic matter to the surface of Earth is affected by energy-dissipation processes such [...] Read more.
A hypothesis in prebiotic chemistry argues that organics were delivered to the early Earth in abundance by meteoritic sources. This study tests that hypothesis by measuring how the transfer of organic matter to the surface of Earth is affected by energy-dissipation processes such as ablation and airbursts. Exogenous delivery has been relied upon as a source of primordial material, but it must stand to reason that other avenues (i.e., hydrothermal vents, electric discharge) played a bigger role in the formation of life as we know it on Earth if exogenous material was unable to deliver significant quantities of organics. For this study, we look at various properties of meteors such as initial velocity and mass of the object, and atmospheric composition to see how meteors with different initial velocities and masses ablate. We find that large meteors do not slow down fast enough and thus impact the surface, vaporizing their components; fast meteors with low masses are vaporized during entry; and meteors with low velocities and high initial masses reach the surface. For those objects that survive to reach the surface, about 60 to >99% of the mass is lost by ablation. Large meteors that fragment are also shown to spread out over increasingly larger areas with increasing mass, and small meteors (~1 mm) are subjected to intense thermal heating, potentially degrading intrinsic organics. These findings are generally true across most atmospheric compositions. These findings provide several caveats to extraterrestrial delivery models that—while a viable point source of organics—likely did not supply as much prebiotic material as an effective endogenous production route. Full article
(This article belongs to the Special Issue Meteorites and the Origin of Life)
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Figure 1
<p>The final mass of a meteorite vs. its initial mass and velocity, under present atmospheric conditions. Normalized mass is the final mass of the object divided by its initial mass.</p>
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<p>As per <a href="#life-08-00013-f001" class="html-fig">Figure 1</a>, for a thin N<sub>2</sub> atmosphere (0.5 atm at the surface).</p>
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<p>As per <a href="#life-08-00013-f001" class="html-fig">Figure 1</a>, with a CO<sub>2</sub> atmosphere (3 atm at the surface).</p>
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<p>Relationship between the total mass of a single meteorite recovered, and the area over which all the fragments were recovered (the strewn field). The regression line has a slope of log A = 1.25 log M − 0.23 (R<sup>2</sup> = 0.47) where A is the radial area and M is the mass. This is not to say that the mass is the direct control on strewn field area, as other factors (angle of entry, meteoroid structural integrity prior to entry) likely play as large of roles in strewn field sizes. Some of this information is not known for these strewn fields as many are meteorite finds. However, that there is a relationship between mass and area is suggestive that the mass controls a portion of the area over which a meteorite is distributed.</p>
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<p>T-t-r (Temperature, time and radius) profile predicted for meteors in contact with a 1000 K surface. This model assumes a total meteor radius of 1 mm, and the dashed lines provide the temperature from the center (0 mm) to the exterior (1 mm) as a function of time.</p>
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<p>The timescale it takes to degrade valine as a function of temperature. The black line is the time required to degrade 50% of the material (a chemical half life), and the red line corresponds to the time required to degrade 99% of the valine.</p>
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<p>Schematic showing that higher velocity objects may vaporize as they travel through the atmosphere, and larger objects vaporize when they impact the surface.</p>
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22 pages, 1845 KiB  
Article
Comet Pond II: Synergistic Intersection of Concentrated Extraterrestrial Materials and Planetary Environments to Form Procreative Darwinian Ponds
by Benton C. Clark and Vera M. Kolb
Life 2018, 8(2), 12; https://doi.org/10.3390/life8020012 - 11 May 2018
Cited by 12 | Viewed by 5918
Abstract
In the “comet pond” model, a rare combination of circumstances enables the entry and landing of pristine organic material onto a planetary surface with the creation of a pond by a soft impact and melting of entrained ices. Formation of the constituents of [...] Read more.
In the “comet pond” model, a rare combination of circumstances enables the entry and landing of pristine organic material onto a planetary surface with the creation of a pond by a soft impact and melting of entrained ices. Formation of the constituents of the comet in the cold interstellar medium and our circumstellar disk results in multiple constituents at disequilibrium which undergo rapid chemical reactions in the warmer, liquid environment. The planetary surface also provides minerals and atmospheric gases which chemically interact with the pond’s organic- and trace-element-rich constituents. Pond physical morphology and the heterogeneities imposed by gravitational forces (bottom sludge; surface scum) and weather result in a highly heterogeneous variety of macro- and microenvironments. Wet/dry, freeze/thaw, and natural chromatography processes further promote certain reaction sequences. Evaporation concentrates organics less volatile than water. Freezing concentrates all soluble organics into a residual liquid phase, including CH3OH, HCN, etc. The pond’s evolutionary processes culminate in the creation of a Macrobiont with the metabolically equivalent capabilities of energy transduction and replication of RNA (or its progenitor informational macromolecule), from which smaller organisms can emerge. Planet-wide dispersal of microorganisms is achieved through wind transport, groundwater, and/or spillover from the pond into surface hydrologic networks. Full article
(This article belongs to the Special Issue Meteorites and the Origin of Life)
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<p>A cometary nucleus or primitive carbonaceous planetoid accomplishes a rare entry event which allows the survival of pristine organic material and ices upon landing on the surface of a planetary body, such as Earth. (Credit: Lockheed Martin).</p>
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<p>Following a relatively low-velocity landing, the ices melt to form a pond with a residual organic-rich central mound as well as bottom sludge, plus liquid with solutes and colloidal particulates, and surface scum. The pond interacts with surface minerals and atmospheric gases to create ideal conditions for diverse prebiotic syntheses. (credit: Lockheed Martin).</p>
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<p>Comet pond phases (<b>a</b>) Schematic of early pond configuration, with satellite pond (s). (<b>b</b>) Gravitational force results in floating scum and sunken sludge (graded bed). (<b>c</b>) In a hot dry climate, evaporation will precipitate salts and form hydrogels. (<b>d</b>) In a cold climate, or at night, ice forms and freeze/thaw cycling can occur. (<b>e</b>) The Macrobiont forms (a consortium of living entities, see text). (<b>f</b>) Escape from the pond can occur via multiple paths, into the air or planetary hydrological network.</p>
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<p>Comet pond phases (<b>a</b>) Schematic of early pond configuration, with satellite pond (s). (<b>b</b>) Gravitational force results in floating scum and sunken sludge (graded bed). (<b>c</b>) In a hot dry climate, evaporation will precipitate salts and form hydrogels. (<b>d</b>) In a cold climate, or at night, ice forms and freeze/thaw cycling can occur. (<b>e</b>) The Macrobiont forms (a consortium of living entities, see text). (<b>f</b>) Escape from the pond can occur via multiple paths, into the air or planetary hydrological network.</p>
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15 pages, 5358 KiB  
Article
Amphiphilic Compounds Assemble into Membranous Vesicles in Hydrothermal Hot Spring Water but Not in Seawater
by Daniel Milshteyn, Bruce Damer, Jeff Havig and David Deamer
Life 2018, 8(2), 11; https://doi.org/10.3390/life8020011 - 10 May 2018
Cited by 60 | Viewed by 9305
Abstract
There is a general assumption that amphiphilic compounds, such as fatty acids, readily form membranous vesicles when dispersed in aqueous phases. However, from earlier studies, it is known that vesicle stability depends strongly on pH, temperature, chain length, ionic concentration and the presence [...] Read more.
There is a general assumption that amphiphilic compounds, such as fatty acids, readily form membranous vesicles when dispersed in aqueous phases. However, from earlier studies, it is known that vesicle stability depends strongly on pH, temperature, chain length, ionic concentration and the presence or absence of divalent cations. To test how robust simple amphiphilic compounds are in terms of their ability to assemble into stable vesicles, we chose to study 10- and 12-carbon monocarboxylic acids and a mixture of the latter with its monoglyceride. These were dispersed in hydrothermal water samples drawn directly from hot springs in Yellowstone National Park at two pH ranges, and the results were compared with sea water under the same conditions. We found that the pure acids could form membranous vesicles in hydrothermal pool water, but that a mixture of dodecanoic acid and glycerol monododecanoate was less temperature-sensitive and assembled into relatively stable membranes at both acidic and alkaline pH ranges. Furthermore, the vesicles were able to encapsulate nucleic acids and pyranine, a fluorescent anionic dye. None of the amphiphiles that were tested formed stable vesicles in sea water because the high ionic concentrations disrupted membrane stability. Full article
(This article belongs to the Special Issue Hydrothermal Vents or Hydrothermal Fields: Challenging Paradigms)
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<p>Acidic clay-lined acid pool (<b>A</b>), silica-rich alkaline Bison Pool (<b>B</b>) and sample vial (<b>C</b>) with Bison Pool water added and shaken to disperse the amphiphilic compounds.</p>
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<p>Decanoic acid (10 carbons) formed droplets at an acidic pH range (Geyser Basin sample) but partially crystallized at the alkaline pH range (Bison Pool sample) even above the transition temperature of 31 °C. All bars show 20 micrometer scales.</p>
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<p>Dodecanoic acid at pH 3.3 (Geyser Basin sample) and pH 7.9 (Bison Pool sample). At both pH ranges the vesicles emerged from melted droplets of the acids.</p>
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<p>Dodecanoic acid (12 carbons), mixed in a 1:1 mole ratio with its monoglyceride, formed stable membranes at both acidic (Geyser Basin) and alkaline (Bison Pool) pH ranges.</p>
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<p>Dodecanoic acid (12 carbons) were mixed in a 1:1 mole ratio with its monoglyceride in pH 7.9 Bison Pool water, then were put through a wet-dry-wet cycle in the presence of 0.1 mM pyranine dye. Large vesicles are barely visible in the phase image (arrows, A) because the very thin bilayer membranes are not clearly resolved. However, the encapsulated pyranine was revealed within vesicles by fluorescence microscopy (B). Some vesicles have excluded the dye for reasons explained in the text.</p>
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<p>Lauric acid–glycerol monolaurate (LA–GML, 10 mM) was mixed with yeast ribosomal RNA in a 2:1 ratio by weight in pH 3.3 Geyser Basin water, then dried on a microscope and rehydrated with the same volume of water. Phase (<b>A</b>) and fluorescence (<b>B</b>) images. (<b>C</b>) A control in which the same lipid mixture was prepared without a wet–dry cycle and put through the gel permeation column in the absence of RNA.</p>
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<p>LA–GML (10 mM) was mixed with lambda DNA in a 2:1 ratio by weight in pH 3.3 Geyser Basin water, then dried on a microscope slide and rehydrated with the same volume of water. Phase (<b>top</b>) and fluorescence (<b>bottom</b>) images.</p>
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<p>Dodecanoic acid (12-carbon lauric acid) in seawater formed crystals protruding from a central mass (<b>A</b>). A mixture of dodecanoic acid and its monoglyceride also formed particulate aggregates surrounded by crystals (<b>B</b>).</p>
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<p>LA–GML put through a wet-dry-wet cycle in seawater formed intractable aggregates (<b>A</b>) that did not encapsulate pyranine dye (<b>B</b>).</p>
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<p>Ionic composition of seawater compared with hydrothermal water from Mammoth Hot Springs in Yellowstone National Park. Data from [<a href="#B16-life-08-00011" class="html-bibr">16</a>].</p>
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19 pages, 2429 KiB  
Review
Mineral Surface-Templated Self-Assembling Systems: Case Studies from Nanoscience and Surface Science towards Origins of Life Research
by Richard J. Gillams and Tony Z. Jia
Life 2018, 8(2), 10; https://doi.org/10.3390/life8020010 - 8 May 2018
Cited by 30 | Viewed by 8171
Abstract
An increasing body of evidence relates the wide range of benefits mineral surfaces offer for the development of early living systems, including adsorption of small molecules from the aqueous phase, formation of monomeric subunits and their subsequent polymerization, and supramolecular assembly of biopolymers [...] Read more.
An increasing body of evidence relates the wide range of benefits mineral surfaces offer for the development of early living systems, including adsorption of small molecules from the aqueous phase, formation of monomeric subunits and their subsequent polymerization, and supramolecular assembly of biopolymers and other biomolecules. Each of these processes was likely a necessary stage in the emergence of life on Earth. Here, we compile evidence that templating and enhancement of prebiotically-relevant self-assembling systems by mineral surfaces offers a route to increased structural, functional, and/or chemical complexity. This increase in complexity could have been achieved by early living systems before the advent of evolvable systems and would not have required the generally energetically unfavorable formation of covalent bonds such as phosphodiester or peptide bonds. In this review we will focus on various case studies of prebiotically-relevant mineral-templated self-assembling systems, including supramolecular assemblies of peptides and nucleic acids, from nanoscience and surface science. These fields contain valuable information that is not yet fully being utilized by the origins of life and astrobiology research communities. Some of the self-assemblies that we present can promote the formation of new mineral surfaces, similar to biomineralization, which can then catalyze more essential prebiotic reactions; this could have resulted in a symbiotic feedback loop by which geology and primitive pre-living systems were closely linked to one another even before life’s origin. We hope that the ideas presented herein will seed some interesting discussions and new collaborations between nanoscience/surface science researchers and origins of life/astrobiology researchers. Full article
(This article belongs to the Special Issue Water–Rock Interactions and Life)
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Graphical abstract

Graphical abstract
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<p>The snapshot time step from molecular dynamics simulations of ethanol layer assembly (into two distinct layers: a thin, ordered monolayer with the thickness of exactly one ethanol molecule, and a second larger less-ordered layer above the monolayer, separated by a gap) on a calcite surface, along with the ethanol spatial density profile associated with it. Reprinted with permission from Pasarín, I. S., Yang, M., Bovet, N., Glyvradal, M., Nielsen, M. M., Bohr, J., Feidenhans’l, R., and Stipp, S. L. S. 2012. “Molecular Ordering of Ethanol at the Calcite Surface.” <span class="html-italic">Langmuir</span> 28 (5):2545–50. [<a href="#B45-life-08-00010" class="html-bibr">45</a>] Copyright 2011 American Chemical Society.</p>
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<p>Atomic force microscope images of a structured DNA microarray self-assembled on a mica (general) surface. Reprinted with permission from Sun, X., Ko, S. H., Zhang, C., Ribbe, A. E., and Mao, C., 2009. “Surface-Mediated DNA Self-Assembly.” <span class="html-italic">J. Am. Chem. Soc.</span> 131 (37):13248-9. [<a href="#B80-life-08-00010" class="html-bibr">80</a>] Copyright 2009 American Chemical Society.</p>
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<p>Atomic force microscope images of the assembly dynamics of SMA (an amyloidogenic light chain variable domain peptide with 114 residues) nanofibrils on a mica surface over time (<b>A</b>–<b>E</b>) show 12–16 h of incubation with image size of 1 µm × 1 µm, while (<b>F</b>) shows 6 days of incubation with image size of 5 µm × 5 µm). It appears that longer incubation times result in longer nanofibrils, such as in (<b>F</b>), as compared to shorter incubation times, such as in (<b>A</b>–<b>E</b>). Reprinted with permission from Zhu, M., Souillac, P. O., Ionescu-Zanetti, C., Carter, S. A., and Fink, A. L. 2002. “Surface-catalyzed Amyloid Fibril Formation.” <span class="html-italic">J. Biol. Chem.</span> 277(52):50914-22. [<a href="#B112-life-08-00010" class="html-bibr">112</a>] Copyright 2002 American Society for Biochemistry and Molecular Biology.</p>
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<p>Transmission electron microscope images of calcium carbonate minerals at different nutrient concentrations and incubation times grown on a self-assembled short peptide nanostructure (Ac-VHVEVS-CONH2), suggesting that mineral growth can be effected by simple peptide self-assemblies. Reprinted with permission from Murai, K., Kinoshita, T., Nagata, K., and Higuchi, M. 2016. “Mineralization of Calcium Carbonate on Multifunctional Peptide Assembly Acting as Mineral Source Supplier and Template.” <span class="html-italic">Langmuir</span> 32 (36):9351–59. [<a href="#B157-life-08-00010" class="html-bibr">157</a>] Copyright 2016 American Chemical Society.</p>
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<p>Synergistic cyclical model of mineral-templated self-assembling systems promoting mineralization. Simple precursors of biomolecules adsorbed onto a mineral surface react and produce biomolecules such as nucleotides or amino acids. Then, some of these adsorbed molecules polymerize into polymers (such as peptides or nucleic acids). Given the optimal conditions and mineral surfaces, biopolymers produced then self-assemble into supramolecular structures such as amyloid fibers or primitive nucleic acid “origamis”. Although not shown in the figure, at any stage in the adsorption, polymerization, and self-assembly pathway (which are all reversible reactions), each molecule can also desorb off of the mineral surface. However, certain self-assembled supramolecular structures (whether on or off the mineral surface) have the ability to promote the formation of new minerals, such as peptide nanostructures catalyzing the formation of hydroxyapatite, similar to modern-day biomineralization processes. The newly produced mineral surfaces could then catalyze and template the synthesis, polymerization, and/or assembly of the same structures that allowed its own mineralization, leading to a synergistic cyclical positive feedback loop by which primitive self-assemblies could have affected the mineral composition of early Earth, and vice-versa.</p>
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15 pages, 16733 KiB  
Article
Cystobasidium alpinum sp. nov. and Rhodosporidiobolus oreadorum sp. nov. from European Cold Environments and Arctic Region
by Benedetta Turchetti, Laura Selbmann, Nina Gunde-Cimerman, Pietro Buzzini, José Paulo Sampaio and Polona Zalar
Life 2018, 8(2), 9; https://doi.org/10.3390/life8020009 - 5 May 2018
Cited by 17 | Viewed by 7020
Abstract
Over 80% of the Earth’s environments are permanently or periodically exposed to temperatures below 5 °C. Cold habitats harbour a wide diversity of psychrophilic and psychrotolerant yeasts. During ecological studies of yeast communities carried out in cold ecosystem in the Italian Alps, Svalbard [...] Read more.
Over 80% of the Earth’s environments are permanently or periodically exposed to temperatures below 5 °C. Cold habitats harbour a wide diversity of psychrophilic and psychrotolerant yeasts. During ecological studies of yeast communities carried out in cold ecosystem in the Italian Alps, Svalbard (Norway, Arctic region), and Portugal, 23 yeast strains that could not be assigned to any known fungal taxa were isolated. In particular, two of them were first identified as Rhodotorula sp., showing the highest degree of D1/D2 sequence identity with Cystobasidum laryngis accounted to only 97% with the type strain (C. laryngis CBS 2221). The other 21 strains, exhibiting identical D1/D2 sequences, had low identity (97%) with Rhodosporidiobolus lusitaniae and Rhodosporidiobolus colostri. Similarly, ITS sequences of the type strains of the most closely related species (93–94%). In a 2-genes multilocus D1/D2 and ITS ML phylogenetic tree, the studied strains pooled in two well separated and supported groups. In order to classify the new 23 isolates based on phylogenetic evidences, we propose the description of two novel species Cystobasidium alpinum sp. nov. and Rhodosporidiobolus oreadorum sp. nov. Full article
(This article belongs to the Special Issue Fungi from Extreme Environments)
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<p>ML 2-genes multilocus phylogeny (ITS and D1/D2) showing the phylogenetic placement of the new taxa described. The tree, based on 42 strains and 1348 nucleotide positions, has been generated using a GTR+IG(4) model calculated using ML in the software MrAIC. Bootstrap values above 80%, calculated from 1000 resampled data sets, are shown.</p>
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<p><span class="html-italic">Cystobasidium alpinum</span> (<b>a</b>) Colonies of DBVPG 10041<sup>T</sup> on MEA after 2 weeks incubation at 15 °C. (<b>b</b>) Polar budding cells of DBVPG 10041<sup>T</sup> on MEA after 1 week incubation at 15 °C.</p>
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<p><span class="html-italic">Rhodosporidium oreadorum</span> Colonies of EXF-3880<sup>T</sup> on PDA (<b>a</b>) and MEA (<b>b</b>) after 1 week of incubation at 20 °C. Polar budding (<b>c</b>) of EXF-3880<sup>T</sup> on MEA after 1 week of incubation at 20 °C. After mating compatible strains on SG agar: mycelium with clamp connections (<b>d</b>), teliospores (<b>e</b>), germinated teliospores with transversely septate basidia and basidiospores (<b>f</b>). Scale bar indicated on figure (<b>c</b>) is also valid for figures (<b>d</b>) and (<b>e</b>).</p>
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