The Chemistry category reflects chemical properties of substances known to be produced or modified by life. Chemical synthesis in biological systems seeks to maximize fitness and adaptive utility. In contrast, abiotic reactions are dictated by kinetics and thermodynamics. This difference can result in diagnostic chemical properties of substances (elements, molecules, compounds, minerals) that serve as proxy for biological or abiotic sources, including the structure of organic molecules, the relative abundance of choice of certain enantiomers, or the isotopic makeup of organic and inorganic compounds. In addition, biological processes can modify the chemical composition of an environment from the cellular to the planetary scale in ways that are distinguishable from abiotic processes — including the composition and relative abundances of elements and molecules within living things (e.g., atmospheric gases, chemical composition of crustal minerals and rocks).
Molecular structure refers to the three-dimensional arrangement of atoms in a substance. Living systems can produce complex molecules that are not observed in abiotic systems. For example, the presence in a sample of a complex polymer with repeating units (monomers), or with a repeating charge (i.e., polyelectrolyte) could be indicative of biological synthesis. Some monomers are also sufficiently complex that their presence alone could point to a biological source, such as the proteinogenic amino acids tryptophan and phenylalanine, or sugar monomers with five or more carbons in their structure.
Living systems utilize only a small subset of monomers (e.g., amino acids, sugars, fatty acids) to synthesize more complex polymers. The relative abundance of each monomer within that subset depends on the structure and functionality of polymers, which are the result of evolutionary adaptations. In contrast, when the same monomers are produced abiotically their relative abundance is a function of their energy of formation, so smaller molecules tend to dominate. The different patterns in the relative abundance of molecules within a certain class of compounds can therefore be used to discriminate biotic sources from abiotic ones (Dorn et al., 2011).
Isomers are compounds with the same formula but a different arrangement of atoms in the molecule, which can confer different properties. There are two general types of isomers: (1) Constitutional (or structural) isomers; and (2) Stereoisomers (which include diastereomers and enantiomers). A hallmark of living systems is their selectivity toward specific isomers involved in biochemical roles. This selectivity can lead to some readily identifiable and measurable molecular qualities that can be used as potential biosignatures (Summons et al., 2008). For example, enantiomers are pairs of the same molecule that are mirror images of each other. Life on Earth often displays a preference toward specific enantiomers (e.g., amino acids; sugars) to build functional polymers (e.g., peptides; nucleic acids), which can result in high levels of enantiomeric excesses in a sample, which might serve as signatures of biological activity (Avnir, 2021). Abiotic organic synthesis tends to form equal mixtures [read more]Isomers are compounds with the same formula but a different arrangement of atoms in the molecule, which can confer different properties. There are two general types of isomers: (1) Constitutional (or structural) isomers; and (2) Stereoisomers (which include diastereomers and enantiomers). A hallmark of living systems is their selectivity toward specific isomers involved in biochemical roles. This selectivity can lead to some readily identifiable and measurable molecular qualities that can be used as potential biosignatures (Summons et al., 2008). For example, enantiomers are pairs of the same molecule that are mirror images of each other. Life on Earth often displays a preference toward specific enantiomers (e.g., amino acids; sugars) to build functional polymers (e.g., peptides; nucleic acids), which can result in high levels of enantiomeric excesses in a sample, which might serve as signatures of biological activity (Avnir, 2021). Abiotic organic synthesis tends to form equal mixtures of enantiomers in the absence of an asymmetric chiral influence (i.e., a physical or chemical process that favors a specific enantiomer), although abiotic organic mixtures, such as those found in carbonaceous chondrites, sometimes display enantiomeric imbalances, which point to abiotic mechanisms of enantioselectivity (e.g., Glavin et al., 2020).
The main elements composing life as we know it are carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur (CHNOPS). Enrichments of these compounds in a sample could be indicative of biological activity, and some of these elements can display remarkably stable ratios both in living systems and the environments they occupy, such as the well-known Redfield ratio of C:N:P=106:16:1. In addition, biological activity can affect the concentrations of trace metals in the environment, and organisms can generate patterns of trace metal abundances that reflect specific aspects of their biochemistry, such as the metallome requirements of methanogenesis (Cameron et al., 2012).
Reactions between chemical species can create patterns in the relative abundances of their stable isotopes that reflect the reaction mechanisms and environmental conditions involved. Isotopes of an element differ in the number of neutrons and therefore in their masses. Because chemical bonds having heavier isotopes have lower vibrational energy states than those with lighter isotopes, bonds having a heavier isotope are stronger (they have larger force constants) than bonds having only lighter isotopes. Molecules with stronger bonds exhibit greater differences in zero-point energies between the molecules’ lighter and heavier isotopes. Compounds having stronger bonds tend to maintain relatively greater abundances of heavier isotopes (e.g., see http://www.soest.hawaii.edu/krubin/GG325/textbook/Chapter09.pdf).
Equilibrium fractionation. Coexisting molecules can achieve mutual chemical and isotopic equilibrium when the forward and backward reactions
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Reactions between chemical species can create patterns in the relative abundances of their stable isotopes that reflect the reaction mechanisms and environmental conditions involved. Isotopes of an element differ in the number of neutrons and therefore in their masses. Because chemical bonds having heavier isotopes have lower vibrational energy states than those with lighter isotopes, bonds having a heavier isotope are stronger (they have larger force constants) than bonds having only lighter isotopes. Molecules with stronger bonds exhibit greater differences in zero-point energies between the molecules’ lighter and heavier isotopes. Compounds having stronger bonds tend to maintain relatively greater abundances of heavier isotopes (e.g., see http://www.soest.hawaii.edu/krubin/GG325/textbook/Chapter09.pdf).
Equilibrium fractionation. Coexisting molecules can achieve mutual chemical and isotopic equilibrium when the forward and backward reactions between them have equal rates, allowing the isotopes to achieve equilibrium distributions among molecules. At chemical equilibrium, molecules having stronger bonds will exhibit greater abundances of heavier isotopes. Differences in isotopic abundances between equilibrated molecules generally decrease at higher temperatures. When isotopic differences between equilibrated molecules can be achieved solely by chemical and physical phenomena, they do not necessarily require biological processes and are not definitive biosignatures. In some cases, enzymatic processes can approach isotopic equilibrium between molecules under conditions where equilibration would otherwise be extremely slow without life (e.g., Kolodny, 1983; Colman 2005), thus evidence of isotopic equilibrium might indicate biological activity.
Kinetic isotope fractionation. A kinetically controlled reaction can arise when its rate is at least somewhat inhibited and the rate from reactant to product is greater than the rate of the reverse reaction. A reactant can still equilibrate with a higher energy (less stable) ‘transition state’ species that ultimately leads to the product. Because the reacting bonds in the ‘transition state’ are typically weaker than the reactant’s bonds, the ‘transition state’ becomes relatively enriched in lighter isotopes. Thus, the product of a kinetically-controlled reaction is typically enriched in lighter isotopes relative to the reactant.
Enzymes can alter reaction kinetics and isotopic discrimination and create distinctive isotopic patterns in biomolecules (e.g., Hayes, 2001). An enzyme’s isotopic discrimination is determined by the molecular structure of its substrate’s ‘transition state’ and the environment within the enzyme’s reaction center. Differences in reaction mechanisms employed by biological versus nonbiological processes can form the basis for distinguishing between isotopic patterns from biotic versus abiotic sources. For example, the biosynthesis of straight-chain lipids involves isotopic discrimination during the formation of 2-carbon subunits that are then linked, creating alternating different 13C/12C values along the lipid’s C backbone (e.g., Monson, 1982). Isotopic differences between groups of biogenic amino acids reflect each group’s biosynthetic pathway and these pathways are highly prevalent in life (e.g., Scott, 2006).
Mass-independent isotopic fractionation. Elements having multiple (>2) stable isotopes can exhibit isotopic patterns that reflect reaction mechanisms and pathways. For example, compounds associated with dissimilatory sulfate reduction, elemental S disproportionation and sulfite disproportionation exhibit patterns of multiple isotopes that can discriminate between these different metabolic processes (Johnston 2005). Different isotopic patterns in mid-ocean ridge hydrothermal deposits indicate abiotic processes (McDermott 2015).
Clumped isotopic patterns. Abundance patterns among four rare isotopologues of methane (13CH4, 12CH3D, 13CH3D and 12CH2D2) can indicate their sources. For example, methane produced by methylotrophic methanogens is more depleted in clumped isotopologues (13CH3D and 12CH2D2) than methane from hydrogenotrophic methanogens (Shuai 2021). These patterns differ from those among methane isotopologues at equilibrium at ambient environmental temperatures.
Parallel observations of associated environments, processes, and geochemical reservoirs are essential (e.g., Des Marais, 2001; Lloyd, 2020). Physicochemical and biological processes transport and transform biologically important elements between their crustal, marine, and atmospheric reservoirs. These processes collectively contribute to the isotopic patterns observed in their abiotic and biotic products. Interpretations of isotope patterns in the geologic record help to decipher our biosphere’s history. Investigating isotopic patterns also promotes an interdisciplinary approach toward determining environmental contexts for interpreting biosignatures (e.g., Chan, 2019).
This taxonomic category is an outlier in that it does not conform to the tier 1 (chemical parameters) and tier 2 (substances) structure. However, it was included in the taxonomic scheme to capture physical/chemical phenomena indicative of compounds that can serve important biochemical or physiological roles (e.g., light absorption, electrochemical properties…). Measurements of those chemical properties can point to the presence of life even if the compounds are unknown (the physical/chemical phenomena can sometimes be measured remotely without the need for compositional analyses). For example, absorption of light within specific regions of the solar spectrum could be indicative of a substance that can capture and convert light energy into chemical energy, akin to biological pigments such as chlorophyll.
This category includes visual observable characteristics, arrangements, and spatial relationships of a visual feature that aid in assessing biogenic / abiogenic origin, irrespective of geochemistry or scale. This content was initially developed from geologic phenomena expected on a rocky planet, though measurement parameters (MPs) should apply to any visually observable target.
Measurement parameters (MPs) include fundamental attributes of biogenic features and abiotic mimics, plus key emergent features of structures (i.e., stromatolites, microbially induced sedimentary structures) that result from microbe-mineral interactions. Emergent features have attributes that individual components of those features do not have. MPs considered here are size distribution, spatial distribution, shape/form, orientation, and texture / fabric of features of interest. These measurable parameters were adapted from classic geological terminologies and designed to be useful and “agnostic” for a visual [read more]This category includes visual observable characteristics, arrangements, and spatial relationships of a visual feature that aid in assessing biogenic / abiogenic origin, irrespective of geochemistry or scale. This content was initially developed from geologic phenomena expected on a rocky planet, though measurement parameters (MPs) should apply to any visually observable target.
Measurement parameters (MPs) include fundamental attributes of biogenic features and abiotic mimics, plus key emergent features of structures (i.e., stromatolites, microbially induced sedimentary structures) that result from microbe-mineral interactions. Emergent features have attributes that individual components of those features do not have. MPs considered here are size distribution, spatial distribution, shape/form, orientation, and texture / fabric of features of interest. These measurable parameters were adapted from classic geological terminologies and designed to be useful and “agnostic” for a visual unknown feature.
The description of structural (visual) attributes included here avoids the use of terms based on (bio)genetic classifications.
Often, companion LDKB entries are included in the “Chemistry” category with a link provided in the relevant “Structure” argument to describe their full value as potential biosignatures.
The distribution of a feature’s dimensions / magnitude.
For example, for laminations, this can refer to laminae thickness distribution, measured in cross-section parallel to its growth direction. For tunnels, this can refer to tunnel diameter distributions and length ranges.
Spatial distribution is the proportion of a 2D surface covered by a feature within an area of interest. Of interest are patterns on surfaces and the spatial frequency (including repetitions) of distinctive features that intersect the surface or its component grains (allochthonous, autochthonous), minerals (detrital, chemical), mineraloids (non-X-ray crystalline), and other components (e.g., dispersed organics, fossils, etc.).
For example, in basalt, tunnels can be distributed along fractures, starting at a void, and providing an opening for water and / or microorganisms to enter the rock.
Shape / form is the geometric shape, morphology, 2D or 3D structure, habit / crystal habit, surface relief, or surface topography.
For example, spherical cells, also called cocci, represent a morphological variety of unicellular organisms and are distinguished from other cellular morphologies, such as rods and filaments. Upon burial and compaction, coccoidal microfossils may undergo deformation to resemble ruptured or deflated spheres (e.g. raisin-like), or may become flattened compressions. Abiotic processes commonly create spherical morphologies, too (i.e. "blueberries" on Mars) and other botryoidal mineral habits.
Orientation is the position of a feature relative to its surroundings and characteristics of the substrate (e.g., “Orientation/lower border of view area”). Orientation includes linear arrangements of features, minerals, and cell-sized objects. Orientation also includes the alignment of structures and their 3D spatial arrangements. One example is tunnels that can be positioned relative to fractures, vesicles (bubbles), minerals, or other tunnels. For example, evidence in support of the biogenic prevalence of the tunnels, includes the density or spacing of microtunnels which is interpreted to reflect sharing of the substrate. An example for filaments includes their directional growth within a mat, which can have a vertical or horizontal orientation relative to the surface, and can be interpreted to reflect phototactic or chemotactic growth.
exture describes the relationships of a rock’s components to one another at the microstructural scale. A rock’s texture records its origin and post-depositional (i.e., taphonomic) history. For a sedimentary rock deposited in the presence of a microbial community, these components include grains, mineral crystals, aqueous precipitates, fossils, organic remains, and other rock fragments. Fabric refers to the spatial arrangement, orientation, and geometric patterns of the visual components, such as grains, filaments, laminae, and any cement of a sedimentary rock that display some level of directionality. A rock fabric can be described by the orientation and pattern produced by single or multiple components of a sediment. For example, stromatolites (laminated geobiological structures) can display distinctive fabrics and textures (such as laminae oriented parallel to the surface of a sedimentary deposit). Fabrics and textures may reveal biogenic attributes and visual patterns that [read more]exture describes the relationships of a rock’s components to one another at the microstructural scale. A rock’s texture records its origin and post-depositional (i.e., taphonomic) history. For a sedimentary rock deposited in the presence of a microbial community, these components include grains, mineral crystals, aqueous precipitates, fossils, organic remains, and other rock fragments. Fabric refers to the spatial arrangement, orientation, and geometric patterns of the visual components, such as grains, filaments, laminae, and any cement of a sedimentary rock that display some level of directionality. A rock fabric can be described by the orientation and pattern produced by single or multiple components of a sediment. For example, stromatolites (laminated geobiological structures) can display distinctive fabrics and textures (such as laminae oriented parallel to the surface of a sedimentary deposit). Fabrics and textures may reveal biogenic attributes and visual patterns that can be diagnostic of various types of microbial input.
The Activity category reflects time-dependent processes that are the result of life’s exchange of matter and energy with the environment. This is manifested in measurable, time-dependent chemical and physical phenomena (e.g., movement of an object). In addition, biological systems can efficiently catalyze specific chemical reactions at rates significantly higher than abiotic processes, and the activity of biological systems can be modulated by environmental factors in ways that are predictable (e.g., seasonal changes), or within well-defined margins (e.g., temperature ranges). The boundaries of the Activity category may appear ambiguous since biological activity can be manifested in the form of chemical compounds or structural features. This category encompasses changes that a substance or structure undergoes over time, but not the substance (Chemistry category) or structure (Structure category) itself.
Reproduction describes the rate of change in numbers of a physical object. The most immediate biological example is cell division, which can occur at rates that are predictable based on physical and chemical conditions (e.g., temperature). Sometimes, biological reproduction results in distinct morphologies such as that observed during different reproductive stages and modes, like binary fission or budding. Measurements of those morphologies would fall under the Structure category, despite their link to reproduction. This highlights the fact that the boundaries between LDMs are not always sharp, and that combinations of LDMs (i.e., change in numbers plus morphology) often increase their individual diagnostic power.
Growth describes the rate of change in size of a physical object. Organisms, or their structural components, typically change in size during a natural life cycle. Populations of organisms (e.g., a biofilm, a forest…) can also change in size at rates that can be predicted based on physical and chemical conditions. Again, the boundaries between growth and reproduction can be fuzzy (in microbiology they are often interchangeable), and for certain organisms, changes in size can be significant during certain stages of the life cycle, also leading to changes in morphology, like in the case of reproduction.
Seasonality describes changes in the properties of an environment in response to cyclic or periodic changes in environmental conditions. For example, seasonal changes in the wavelength-dependent reflectance of a planetary surface or in its atmospheric composition as a function of the periodic variation in the distance to its star or the latitude of its subsolar point (e.g., Olson et al., 2018). Measurements of seasonality (and growth) are often applied to the search for evidence of life at planetary scales (e.g., exoplanets) by means of remote sensing methods.
Motion describes the change in position of a physical object. The changes in direction of motile organisms appear purposeful and are inconsistent with drift or Brownian motion (Nadeau et al., 2018). Biological motion can be interrupted by changes in temperature, radiation or addition of chemical compounds, and certain organisms navigate in response to various physicochemical parameters (taxis) including light, magnetic fields, or chemical gradients, which are testable means to differentiate biological activity from abiotic phenomena.
Catalysis describes a hastening of chemical reactions leading to a lesser amount of time needed to reach chemical equilibrium between reactants and products at given environmental conditions. For example, reduction of CO2 to CH4, even if the latter is the thermodynamically stable species, can take billions of years in the absence of life (e.g., Etiope and Sherwood Lollar, 2013), but is much faster if living organisms deriving energy from this reaction use enzymes to hasten its completion.
