On the early stages of the evolution of the geosphere and biosphere

The conditions necessary for the existence of nucleic-protein life are as follows: the presence of liquid water, an atmosphere, and a magnetic field (all of which protect from meteorites, abrupt changes in temperature, and a flow of charged particles from space) and the availability of nutrients (macro-and microelements in the form of dissolved compounds). In the evolution of the geosphere, complex interference of irreversible processes (general cooling, gravitational differentiation of the Earth’s interior, dissipation of hydrogen, etc.) with cyclic processes of varying natures and periodicities (from the endogenic cycles “from Pangea to Pangea” to Milankovitch cycles), these conditions have repeatedly changed; hence, in the coevolution of the geosphere and biosphere, the vector of irreversible evolution was determined by the geosphere. Only with the appearance of the ocean as a global system of homeostasis, which provided the maintenance and leveling of nutrient concentrations in the hydrosphere, and the conveyor of nutrients from the mantle, “the film of life” could begin its expansion from the source of the nutrients. Life itself is a system of homeostasis, but not due to the global size and a vast buffer capacity, but because of the high rate of reactions and presence of a program (genome) that allowed its development (ontogeny) independent from the outside environment. The early stages of the origin and evolution of the biosphere (from the RNA-world to the development of the prokaryotic ecosystems) were characterized by the domination of chemotrophic ecosystems. The geographical ranges of these ecosystems were directly or indirectly (through the atmosphere and hydrosphere) tied to the sources of nutrients in the geosphere, which were in turn connected to various sources of volcanic and geotectonic activity (geothermal waters, “black smokers” along the rift zones, etc.). This gave the biosphere consisting of chemotrophic ecosystems a mosaic appearance composed of separate local oases of life. The decrease of methane and accumulation of O₂ in the atmosphere in the geological evolution of the Earth caused the extinction of chemotrophic ecosystems and directed evolution of the biosphere toward autotrophy. Autotrophic photosynthesis gave the biosphere an energy source that was not connected to the geosphere, and for the first time allowed its liberation from the geosphere by developing its own vector of evolution. This vector resulted in the biosphere forming a continuous film of life on the planet by capturing the continents and occupying pelagic and abyssal zones, and the appearance of eukaryotes. The geosphere formed biogeochemical cycles in parallel to the geochemical ones, and comparable in the annual balances of participating matter.     

Evolution of geological processes on the early earth and their impact on the early biosphere

Although life already existed in the Paleoarchean (ca. 3.8 Ga), rapid development of the biosphere began only in the Paleoproterozoic, from 2.4–2.3 Ga; this eventually resulted in the emergence of multicellular organisms. This event coincided in time with the irreversible replacement of tectonomagmatic activity, when high-Mg magmatism of the Early Precambrian, derived from depleted mantle, was irreversibly replaced by geochemically enriched Fe-Ti basalts similar to Phanerozoic intraplate magmas. The new type of magmas was characterized by the increased and high content of Fe, Ti, Cu, P, Mn, alkalis, LREE, and other incompatible elements (Zr, Ba, Sr, U, Th, F, etc.), which are required for metabolism and fermentation. It is proposed that this event promoted changes in ecological conditions and rapid development of the biosphere, providing the Earth’s surface with qualitatively new biotic material.     

Life conditions on the early Earth after 4.0 Ga

The organization level of Precambrian fossils is the most reliable indicator of the state and parameters of the biosphere, such as the atmosphere composition, average temperature of the earth’s surface, and others. At present, cyanobacteria, unicellular and multicellular eukaryotes, and coelomates are considered to appear in the geological history of the Earth much earlier than it was supposed previously. Our knowledge and ideas of the early Earth are very important for considering the problems of the origin of life. A key boundary of the earliest period was probably about 4 Ga. This boundary is between the periods documented and undocumented by the geological record. The Earth history and probable surface conditions before 4 Ga are considered by L.M. Mukhina, A.V. Vityazeva, G.V. Pechernikova, and L.V. Ksanfomaliti in this volume.     

Secular oscillations in the stratigraphic record—an acute debate

Work on secular oscillations of the ocean-atmosphere system has reached a particularly interesting stage, characterized by competing models that alternately invoke endogenic and celestial drivers. The most widely accepted concept is the icehouse-greenhouse cycle with an approximate length of 300 Ma. The concept holds that in the past 600 Ma the Earth has oscillated between a glacial mode and one of rather warm, equable climate; correlative changes in mineralogy of marine carbonates and evaporites as well as abundance of igneous intrusions point to an endogenic cause, most probably variations in the rates of plate motion and mantle convection that modulate sea level and the level of atmospheric CO₂ and thus climate. However, the 300-Ma icehouse-greenhouse cycle leaves important data unexplained. The history of glaciations as well as certain climate indicators, such as sea-surface temperatures derived from oxygen isotopes, indicate oscillations with wave lengths of 130–150 Ma. Such oscillations have been observed in the reconstructed cosmic-ray flux of the past 1000 Ma; they may modulate climate via low-level cloud cover. The debate about long-term climate oscillations poses several distinct questions for sedimentary geologists.     

Atmospheric composition and climate on the early Earth

Oxygen isotope data from ancient sedimentary rocks appear to suggest that the early Earth was significantly warmer than today, with estimates of surface temperatures between 45 and 85 degrees C. We argue, following others, that this interpretation is incorrect-the same data can be explained via a change in isotopic composition of seawater with time. These changes in the isotopic composition could result from an increase in mean depth of the mid-ocean ridges caused by a decrease in geothermal heat flow with time. All this implies that the early Earth was warm, not hot.A more temperate early Earth is also easier to reconcile with the long-term glacial record. However, what triggered these early glaciations is still under debate. The Paleoproterozoic glaciations at approximately 2.4Ga were probably caused by the rise of atmospheric O2 and a concomitant decrease in greenhouse warming by CH4. Glaciation might have occurred in the Mid-Archaean as well, at approximately 2.9Ga, perhaps as a consequence of anti-greenhouse cooling by hydrocarbon haze. Both glaciations are linked to decreases in the magnitude of mass-independent sulphur isotope fractionation in ancient rocks. Studying both the oxygen and sulphur isotopic records has thus proved useful in probing the composition of the early atmosphere.     

Humic substances in the early biosphere

Humic substances and their organic-mineral compounds are the first stage in transformation of biotic residues into stable geopolymers, which comprise the main reservoir of organic C in the biosphere. Early appearance of HS in the Earth’s history is of principal importance for the understanding of geo-biological processes on land in the past. However, there is no fossil record of HS before land colonization by lignified vegetation (400 Ma). When the first soil HS were formed? Could HS be synthesized in the Precambrian before plants and mosses terrestrialization? The formation of HS occurs in mesophilic aerobic conditions and requires presence of oxidative catalysts and production of aromatic (phenolic) precursors by biota. In this paper, humification processes in algo-myco-bacterial and lichen communities are discussed from actualistic point of view. These communities are considered as a relict ecosystem, which could dominate on land during the Neoproterozoic-Early Paleozoic (1–0.5 Ga).     

Mars Primordial Crust: Unique Sites for Investigating Proto-biologic Properties

The Martian meteorite collection suggests that intact outcrops or boulder-scale fragments of the 4.5 Ga Martian crust exist within tens of meters of the present day surface of Mars. Mars may be the only planet where such primordial crust samples, representing the first 100 Ma of a planet’s environment, are available. The primordial crust has been destroyed on Earth by plate tectonics and other geological phenomena and is buried on the Moon under hundreds or thousands of meters of megaregoltih. Early Mars appears to have been remarkably similar to early Earth, and samples of rock from the first few Ma or first 100 Ma may reveal “missing link” proto-biological forms that could shed light on the transition from abiotic organic chemistry to living cells. Such organic snapshots of nascent life are unlikely to be found on Earth. Presented at: National Workshop on Astrobiology: Search for Life in the Solar System, Capri, Italy, 26 to 28 October, 2005.     

Borate Minerals and Origin of the RNA World

The RNA World is generally thought to have been an important link between purely prebiotic (>3.7 Ga) chemistry and modern DNA/protein biochemistry. One concern about the RNA World hypothesis is the geochemical stability of ribose, the sugar moiety of RNA. Prebiotic stabilization of ribose by solutions associated with borate minerals, notably colemanite, ulexite, and kernite, has been proposed as one resolution to this difficulty. However, a critical unresolved issue is whether borate minerals existed in sufficient quantities on the primitive Earth, especially in the period when prebiotic synthesis processes leading to RNA took place. Although the oldest reported colemanite and ulexite are 330 Ma, and the oldest reported kernite, 19 Ma, boron isotope data and geologic context are consistent with an evaporitic borate precursor to 2400-2100 Ma borate deposits in the Liaoning and Jilin Provinces, China, as well as to tourmaline-group minerals at 3300–3450 Ma in the Barberton belt, South Africa. The oldest boron minerals for which the age of crystallization could be determined are the metamorphic tourmaline species schorl and dravite in the Isua complex (metamorphism between ca. 3650 and ca. 3600 Ma). Whether borates such as colemanite, ulexite and kernite were present in the Hadean (>4000 Ma) at the critical juncture when prebiotic molecules such as ribose required stabilization depends on whether a granitic continental crust had yet differentiated, because in its absence we see no means for boron to be sufficiently concentrated for borates to be precipitated.     

Evolutionary timing of the origins of mesophilic sulphate reduction and oxygenic photosynthesis: a phylogenomic dating approach

Until recently, the deep‐branching relationships in the bacterial domain have been unresolved. A new phylogenetic approach (termed compartmentalization) was able to resolve these deep‐branching relationships successfully by using a large number of genes from whole genome sequences and by reducing long branch attraction artefacts. This new, well‐resolved phylogenetic tree reveals the evolutionary relationships between diverse bacterial groups that leave important traces in the geological record. It shows that mesophilic sulphate reducers originated before the Cyanobacteria, followed by the origination of sulphur‐ and pyrite‐oxidizing bacteria after oxygen became available in the biosphere. This evolutionary pattern mirrors a similar pattern in the Palaeoproterozoic geological record. Sulphur isotopic fractionation records indicate that large‐scale bacterial sulphate reduction began in marine environments around 2.45 billion years ago (Ga), followed by rapid oxygenation of the atmosphere about 2.3 or 2.2 Ga. Oxygenation was then followed by increasing oceanic sulphate concentrations (probably owing to pyrite oxidation and continental weathering), which then resulted in the disappearance of banded iron formations by 1.8 Ga. The similarity between the phylogenetic and geological records suggests that the geochemical changes observed on the Palaeoproterozoic Earth were caused by major origination events in the mesophilic bacteria, and that these geochemical changes then caused additional origination events, such as aerobic respiration. If so, then constraints on divergence dates can be established for many microbial groups, including the Cyanobacteria, mesophilic bacteria, mesophilic sulphate reducers, methanotrophs, several anoxygenic phototrophs, as well as for mitochondrial endosymbiosis. These dates may also help to explain a large number of other changes in the geological record of the Neoarchean and Palaeoproterozoic Earth. This hypothesis, however, does not agree with the finding of cyanobacterial and eukaryote lipids at 2.7 Ga, and suggests that further work needs to be done to elucidate the discrepancies in both these areas.     

Changes in terrestrial biota before the Permian-Triassic ecological crisis

The period around the Permian-Triassic boundary was marked by one of the most important and interesting events in the evolution of life. The diversity of both marine and continental biotas decreased. The changes were global and led to the establishment of the new Mesozoic World. Transformations of the organic world constituted a single process with changes in the inorganic components of the biosphere. The preceding glacial period had ended and the “cool,” zonal, and markedly seasonal climate was replaced by a “warm,” “equable,” virtually non-seasonal and azonal climate. The new climatic organization remained on Earth for more than two hundred million years. The biotic crisis was global: it involved the sea, the land, and inland waters. The changes on land began earlier and more superficial. The principal events were in the Kazanian and Vyatkian, before the end of the Permian. The crisis was caused to a greater extent by biospheric processes than by momentary external influences, the latter at most triggering the crisis.     

A second species of the squid genus <Emphasis Type="Italic">Kondakovia</Emphasis> (Cephalopoda: Onychoteuthidae) from the sub-Antarctic

A new species of squid, Kondakovia nigmatullini, is described based on two specimens from sub-Antarctic waters of the Southwest Atlantic. The new species is characterised by short, broad, rhombic fins, long rostrum of the gladius (~13% ML); very weak development or absence of longitudinal ridges on the mantle surface; some reticulate folds on mantle surface; 23–27 club hooks; subequal funnel/mantle components of the locking apparatus, and a tricuspid rachidian, with small lateral cusps.     

Animal survival strategies in Neoproterozoic ice worlds

The timing of the first appearance of animals is of crucial importance for understanding the evolution of life on Earth. Although the fossil record places the earliest metazoans at 572–602 Ma, molecular clock studies suggest a far earlier origination, as far back as ~850 Ma. The difference in these dates would place the rise of animal life into a time period punctuated by multiple colossal, potentially global, glacial events. Although the two schools of thought debate the limitations of each other's methods, little time has been dedicated to how animal life might have survived if it did arise before or during these global glacial periods. The history of recent polar biota shows that organisms have found ways of persisting on and around the ice of the Antarctic continent throughout the Last Glacial Maximum (33–14 Ka), with some endemic species present before the breakup of Gondwana (180–23 Ma). Here we discuss the survival strategies and habitats of modern polar marine organisms in environments analogous to those that could have existed during Neoproterozoic glaciations. We discuss how, despite the apparent harshness of many ice covered, sub-zero, Antarctic marine habitats, animal life thrives on, in and under the ice. Ice dominated systems and processes make some local environments more habitable through water circulation, oxygenation, terrigenous nutrient input and novel habitats. We consider how the physical conditions of Neoproterozoic glaciations would likely have dramatically impacted conditions for potential life in the shallows and erased any possible fossil evidence from the continental shelves. The recent glacial cycle has driven the evolution of Antarctica's unique fauna by acting as a “diversity pump,” and the same could be true for the late Proterozoic and the evolution of animal life on Earth, and the existence of life elsewhere in the universe on icy worlds or moons.     

Geological constraints on the origin of oxygenic photosynthesis

This article examines the geological evidence for the rise of atmospheric oxygen and the origin of oxygenic photosynthesis. The evidence for the rise of atmospheric oxygen places a minimum time constraint before which oxygenic photosynthesis must have developed, and was subsequently established as the primary control on the atmospheric oxygen level. The geological evidence places the global rise of atmospheric oxygen, termed the Great Oxidation Event (GOE), between ~2.45 and ~2.32 Ga, and it is captured within the Duitschland Formation, which shows a transition from mass-independent to mass-dependent sulfur isotope fractionation. The rise of atmospheric oxygen during this interval is closely associated with a number of environmental changes, such as glaciations and intense continental weathering, and led to dramatic changes in the oxidation state of the ocean and the seawater inventory of transition elements. There are other features of the geologic record predating the GOE by as much as 200–300 million years, perhaps extending as far back as the Mesoarchean–Neoarchean boundary at 2.8 Ga, that suggest the presence of low level, transient or local, oxygenation. If verified, these features would not only imply an earlier origin for oxygenic photosynthesis, but also require a mechanism to decouple oxygen production from oxidation of Earth’s surface environments. Most hypotheses for the GOE suggest that oxygen production by oxygenic photosynthesis is a precondition for the rise of oxygen, but that a synchronous change in atmospheric oxygen level is not required by the onset of this oxygen source. The potential lag-time in the response of Earth surface environments is related to the way that oxygen sinks, such as reduced Fe and sulfur compounds, respond to oxygen production. Changes in oxygen level imply an imbalance in the sources and sinks for oxygen. Changes in the cycling of oxygen have occurred at various times before and after the GOE, and do not appear to require corresponding changes in the intensity of oxygenic photosynthesis. The available geological constraints for these changes do not, however, disallow a direct role for this metabolism. The geological evidence for early oxygen and hypotheses for the controls on oxygen level are the basis for the interpretation of photosynthetic oxygen production as examined in this review.     

The origin and early evolution of plants

Plant (archaeplastid) evolution has transformed the biosphere, but we are only now beginning to learn how this took place through comparative genomics, phylogenetics, and the fossil record. This has illuminated the phylogeny of Archaeplastida, Viridiplantae, and Streptophyta, and has resolved the evolution of key characters, genes, and genomes – revealing that many key innovations evolved long before the clades with which they have been casually associated. Molecular clock analyses estimate that Streptophyta and Viridiplantae emerged in the late Mesoproterozoic to late Neoproterozoic, whereas Archaeplastida emerged in the late-mid Palaeoproterozoic. Together, these insights inform on the coevolution of plants and the Earth system that transformed ecology and global biogeochemical cycles, increased weathering, and precipitated snowball Earth events, during which they would have been key to oxygen production and net primary productivity (NPP).     

Biogeochemical transformations after the emergence of oxygenic photosynthesis and conditions for the first rise of atmospheric oxygen

The advent of oxygenic photosynthesis represents the most prominent biological innovation in the evolutionary history of the Earth. The exact timing of the evolution of oxygenic photoautotrophic bacteria remains elusive, yet these bacteria profoundly altered the redox state of the ocean–atmosphere–biosphere system, ultimately causing the first major rise in atmospheric oxygen (O₂)—the so-called Great Oxidation Event (GOE)—during the Paleoproterozoic (~2.5–2.2 Ga). However, it remains unclear how the coupled atmosphere–marine biosphere system behaved after the emergence of oxygenic photoautotrophs (OP), affected global biogeochemical cycles, and led to the GOE. Here, we employ a coupled atmospheric photochemistry and marine microbial ecosystem model to comprehensively explore the intimate links between the atmosphere and marine biosphere driven by the expansion of OP, and the biogeochemical conditions of the GOE. When the primary productivity of OP sufficiently increases in the ocean, OP suppresses the activity of the anaerobic microbial ecosystem by reducing the availability of electron donors (H₂ and CO) in the biosphere and causes climate cooling by reducing the level of atmospheric methane (CH₄). This can be attributed to the supply of OH radicals from biogenic O₂, which is a primary sink of biogenic CH₄ and electron donors in the atmosphere. Our typical result also demonstrates that the GOE is triggered when the net primary production of OP exceeds >~5% of the present oceanic value. A globally frozen snowball Earth event could be triggered if the atmospheric CO₂ level was sufficiently small (<~40 present atmospheric level; PAL) because the concentration of CH₄ in the atmosphere would decrease faster than the climate mitigation by the carbonate–silicate geochemical cycle. These results support a prolonged anoxic atmosphere after the emergence of OP during the Archean and the occurrence of the GOE and snowball Earth event during the Paleoproterozoic.     

Dating phototrophic microbial lineages with reticulate gene histories

Phototrophic bacteria are among the most biogeochemically significant organisms on Earth and are physiologically related through the use of reaction centers to collect photons for energy metabolism. However, the major phototrophic lineages are not closely related to one another in bacterial phylogeny, and the origins of their respective photosynthetic machinery remain obscured by time and low sequence similarity. To better understand the co‐evolution of Cyanobacteria and other ancient anoxygenic phototrophic lineages with respect to geologic time, we designed and implemented a variety of molecular clocks that use horizontal gene transfer (HGT) as additional, relative constraints. These HGT constraints improve the precision of phototroph divergence date estimates and indicate that stem green non‐sulfur bacteria are likely the oldest phototrophic lineage. Concurrently, crown Cyanobacteria age estimates ranged from 2.2 Ga to 2.7 Ga, with stem Cyanobacteria diverging ~2.8 Ga. These estimates provide a several hundred Ma window for oxygenic photosynthesis to evolve prior to the Great Oxidation Event (GOE) ~2.3 Ga. In all models, crown green sulfur bacteria diversify after the loss of the banded iron formations from the sedimentary record (~1.8 Ga) and may indicate the expansion of the lineage into a new ecological niche following the GOE. Our date estimates also provide a timeline to investigate the temporal feasibility of different photosystem HGT events between phototrophic lineages. Using this approach, we infer that stem Cyanobacteria are unlikely to be the recipient of an HGT of photosystem I proteins from green sulfur bacteria but could still have been either the HGT donor or the recipient of photosystem II proteins with green non‐sulfur bacteria, prior to the GOE. Together, these results indicate that HGT‐constrained molecular clocks are useful tools for the evaluation of various geological and evolutionary hypotheses, using the evolutionary histories of both genes and organismal lineages.     

The geobiological nitrogen cycle: From microbes to the mantle

Nitrogen forms an integral part of the main building blocks of life, including DNA, RNA, and proteins. N₂ is the dominant gas in Earth's atmosphere, and nitrogen is stored in all of Earth's geological reservoirs, including the crust, the mantle, and the core. As such, nitrogen geochemistry is fundamental to the evolution of planet Earth and the life it supports. Despite the importance of nitrogen in the Earth system, large gaps remain in our knowledge of how the surface and deep nitrogen cycles have evolved over geologic time. Here, we discuss the current understanding (or lack thereof) for how the unique interaction of biological innovation, geodynamics, and mantle petrology has acted to regulate Earth's nitrogen cycle over geologic timescales. In particular, we explore how temporal variations in the external (biosphere and atmosphere) and internal (crust and mantle) nitrogen cycles could have regulated atmospheric pN₂. We consider three potential scenarios for the evolution of the geobiological nitrogen cycle over Earth's history: two in which atmospheric pN₂ has changed unidirectionally (increased or decreased) over geologic time and one in which pN₂ could have taken a dramatic deflection following the Great Oxidation Event. It is impossible to discriminate between these scenarios with the currently available models and datasets. However, we are optimistic that this problem can be solved, following a sustained, open‐minded, and multidisciplinary effort between surface and deep Earth communities.     

Terre et Vie : des histoires imbriquées

The Earth's history consists in recurrent flashbacks of similar events and scenarios: redistribution of continents, orogenic cycles, glaciations, marine transgressions and regressions, etc. In contrast, Life's history evolves according to a succession of stages: prokaryotic stage, eukaryotic cell stage, pluricellular organism stage, terrestrialization, development of animal societies, hominization. With each successive stage the biosphere rises to a higher level of organization and complexity. This evolution results from the natural trend of living organisms to extend their control over the entire planet while they progressively escape the constraints of the aquatic environments and climates. During the last four billion years close and complex interactions prevailed between the history of both the Earth and Life. Living organisms have a profound effect on their environment and on the processes of the Earth dynamics, while the planetary environment controls the evolution of living species. Nevertheless, from time to time, the fragile equilibrium established between Earth's and Life's dynamics breaks down and triggers mass extinctions. It is presently the case of the increasing impact of human activities on the integrity of our planet, a major challenge for humankind during the 21st century.     

Maximum entropy production and general trends in biospheric evolution

The biosphere has greatly shaped the past evolution of the Earth system. Here I argue that life evolved to maximize planetary entropy production. The evolution of the Earth system through time has thus evolved as far away from thermodynamic equilibrium as possible. I describe the implications of this hypothesis for the evolution of the global cycles of water and carbon and the implied consequences for biospheric evolution. This thermodynamic perspective of Earth’s biospheric evolution extends the views of Vernadski and Lovelock and puts it on a quantitative foundation.