Ecosphere,biosphere, or Gaia? What to call the global ecosystem

The terms biosphere, ecosphere, and Gaia are used as names for the global ecosystem. However, each has more than one meaning. Biosphere can mean the totality of living things residing on the Earth, the space occupied by living things, or life and life-support systems (atmosphere, hydrosphere, lithosphere, and pedosphere). Ecosphere is used as a synonym of biosphere and as a term for zones in the universe where life as we know it should be sustainable. Gaia is similar to biosphere (in the sense of life and life-support systems) and ecosphere (in the sense of biosphere as life and life-support systems), but, in its most extreme form, refers to the entire planet as a living entity. A case is made for avoiding the term Gaia (at least as a name for the planetary ecosystem), restricting biosphere to the totality of living things, and adopting the ecosphere as the most apt name for the global ecosystem.     

Gaia again

The ideas of the Gaia hypothesis from the 1960s are today largely included in global ecology and Earth system sciences. The interdependence between biosphere, oceans, atmosphere and geosphere is well-established by data from global monitoring. Nevertheless the theory underlying the holistic view of the homeostatic Earth has remained obscure. Here the foundations of Gaia theory are examined from the recent formulation of the 2nd law of thermodynamics as an equation of motion. According to the principle of increasing entropy, all natural processes, inanimate just as animate, consume free energy, the thermodynamic driving force. All species, abiotic just as biotic are viewed as mechanisms of energy transduction for the global system to evolve toward a stationary state in its surroundings. The maximum entropy state displays homeostasis by being stable against internal fluctuations. When surrounding conditions change or when new mechanisms emerge, the global system readjusts its flows of energy to level newly appeared gradients. Thus, the propositions of Gaia theory and holistic understanding of the global system are recognized as consequences of thermodynamic imperatives.     

Natural selection through survival alone,and the possibility of Gaia

Here I advance two related evolutionary propositions. (1) Natural selection is most often considered to require competition between reproducing “individuals”, sometimes quite broadly conceived, as in cases of clonal, species or multispecies-community selection. But differential survival of non-competing and non-reproducing individuals will also result in increasing frequencies of survival-promoting “adaptations” among survivors, and thus is also a kind of natural selection. (2) Darwinists have challenged the view that the Earth’s biosphere is an evolved global homeostatic system. Since there is only one biosphere, reproductive competition cannot have been involved in selection for such survival-promoting adaptations, they claim. But natural selection through survival could reconcile Gaia with evolutionary theory.     

Open systems living in a closed biosphere: a new paradox for the Gaia debate

While energetically open, the biosphere is appreciably closed from the standpoint of matter exchange. Matter cycling and recycling is hence a necessary and emergent property of the global-scale system known as Gaia. But how can an aggregate of open-system life forms have evolved and persisted for billions of years within a planetary system that is largely closed to matter influx and outflow? The puzzling nature of a closed yet persistent biosphere draws our attention to the course of evolution of fundamental metabolic strategies and matter-capture techniques. It suggests a facet of the Gaia hypothesis, framed in terms of persistence. The oceans, atmosphere, soils and biota constitute a complex system which maintains and adjusts matter cycling and recycling within the constraints of planetary closure such that open-system forms of life can persist. This weaker version of the Gaia hypothesis may be useful because it readily lends itself to at least one form of test. What is the solution to the closed biosphere puzzle, and does it indicate that Gaia merits status as a discrete entity? We suggest several disciplines within the field of biology that might provide tools and perspectives toward reaching a solution. These disciplines include artificial closed ecosystems, prokaryote evolution, the nexus of thermodynamics and evolutionary biology, and hierarchy theory in ecosystem modeling and evolution theory.     

Do evolution and ecology need the Gaia hypothesis?

Gaia theory, which describes the life-environment system of the Earth as stable and self-regulating, has remained at the fringes of mainstream biological science owing to its historically inadequate definition and apparent incompatibility with individual-level natural selection. The key issue is whether and why the biosphere might tend towards stability and self-regulation. We review the various ways in which these issues have been addressed by evolutionary and ecological theory, and relate these to 'Gaia theory'. We then ask how this theory extends the perspectives offered by these disciplines, and how it might be tested by novel modelling approaches and laboratory experiments using emergent technologies.     

Gaia as a complex adaptive system

We define the Gaia system of life and its environment on Earth, review the status of the Gaia theory, introduce potentially relevant concepts from complexity theory, then try to apply them to Gaia. We consider whether Gaia is a complex adaptive system (CAS) in terms of its behaviour and suggest that the system is self-organizing but does not reside in a critical state. Gaia has supported abundant life for most of the last 3.8 Gyr. Large perturbations have occasionally suppressed life but the system has always recovered without losing the capacity for large-scale free energy capture and recycling of essential elements. To illustrate how complexity theory can help us understand the emergence of planetary-scale order, we present a simple cellular automata (CA) model of the imaginary planet Daisyworld. This exhibits emergent self-regulation as a consequence of feedback coupling between life and its environment. Local spatial interaction, which was absent from the original model, can destabilize the system by generating bifurcation regimes. Variation and natural selection tend to remove this instability. With mutation in the model system, it exhibits self-organizing adaptive behaviour in its response to forcing. We close by suggesting how artificial life ('Alife') techniques may enable more comprehensive feasibility tests of Gaia.     

Deep-sea research ground for the study of living matter properties in extreme conditions

The Black Sea hollow bottom is a promising research ground in the field of deep-sea radiochemoecology and exobiology. It has turned out to be at the intersection of the earth and cosmic scientific interests such as deep-sea marine radiochemoecology from the perspective of the study of extreme biogeocenological properties of the Earth biosphere and exobiology from the standpoint of the study of life phenomena (living matter) outside the Earth biosphere, i.e. on other planets and during hypothetical transfer of spores in the outer space. The potential of this ground is substantiated with the data published by the author and co-workers on accumulation of 90Sr, 137Cs and Pu isotopes with silts of bathyal pelo-contour, on the quality of deep-sea hydrogen sulphide waters (after their contact with air) for vital functions of planktonic and benthic aerobes, as well as the species composition of marine, freshwater and terrestrial plants grown from the spores collected from the bottom sediments of the Black Sea bathyal. Discussion was based on V.I. Vernadsky's ideas about the living matter and biosphere, which allowed conclusions about the biospheric and outer space role of the described phenomena.     

The fundamental processes in ecology: a thought experiment on extraterrestrial biospheres

Ecological science is often organised as a hierarchical series of entities: genes, individuals, populations, species, communities, ecosystems and biosphere. Here, I consider an alternative process‐based approach to ecology, and analyse the nature of the fundamental processes in ecology. These fundamental processes are discussed in the context of the following question:‘for any planet with carbon‐based life, which persists over geological time scales, what are the minimum set of ecological processes that must be present?‘I suggest that the following processes would be present on any such planet: energy flow, multiple guilds, ecological tradeo ffs leading to within‐guild biodiversity, ecological hypercycles, merging of organismal and ecological physiology, carbon sequestration and possibly photosynthesis. Nutrient cycling is described as an emergent property of these fundamental processes. I discuss reasons why a biosphere based on a single species with no nutrient cycling is very unlikely to exist. I also describe the concept of ‘Gaian effect’. This suggests that some processes will always tend to extend the lifespan of a biosphere in which they develop (positive Gaian effect) while others could either increase or decrease (negative Gaian effect) such a lifespan. These ideas are discussed in the context of astrobiology, ecosystem services, conservation biology and Gaia theory.     

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.     

Molten earth and the origin of prebiological molecules.

Evidence for the molten Earth at its accretion time has been accumulated through the geochemical investigations and the observations of the surfaces of planets by space probes such as Venera 8, Mariner 9, Surveyor, Luna, and Apollo. The primitive terrestrial atmosphere might have been derived from the volcanic gases, as suggested by Rubey, but of a higher temperature than so far assumed. A thermochemical calculation of the composition of the volcanic gas suggests the following possibilities: (1) Large amounts of H2 and CO were present in the primitive atmosphere. This gives a theoretical basis for the HCN-production experiment by Abelson. (2) HCHO and NH3 existed in the primitive oceans, of the amount comparable with the weight of the present biosphere. (3) Plenty of NO3-, SO4, and PO4 were expected in the primitive oceans. The NO3- ions might have been useful for the nitrate respiration advocated by Egami. In an appendix, it is argued, on ;he basis of the observational evidence of the exospheric temperatures of planets by space probes, that a highly reducing atmosphere would (if it existed on the primitive Earth) have disappeared very quickly due to the thermal escape of hydrogen from its exosphere.     

盖娅假说:在争议中发展

自从1972年Lovelock提出盖娅假说已经过去了40年,但围绕它的争议却从未停止过。盖娅假说在反对者的批评中与支持者的证明中不断发展。当前,最极端形式的盖娅假说基本上已被摒弃,尤其是那种明显带有目的论的说法。弱盖娅提出的"有机体可以影响他们的环境,有机体与环境的反馈耦合可以塑造两者的进化"这两个观点也已经是普遍接受的事实。除此之外,盖娅假说提出的其他3个命题却饱受争议。(1)内在平衡的盖娅:生物调节反馈有助于环境的内在平衡。反对者认为,生物反馈稳定全球环境的说法,与冰芯记录和大量的气候反馈研究结果相矛盾的。支持者认为,地球生物-环境系统的内在平衡可以产生于正负反馈的混合。盖娅假说关心的是地球几十亿年的历史,盖娅假说在较短时间尺度内可证伪,并不意味着其在较长时间尺度内也可证伪。(2)最优的盖娅:生物调节环境,使环境更加适合生物的生存。关于有机体的繁荣主要是由于他们对环境的改变,还是由于他们对环境的适应,目前尚未有结论。但盖娅的支持者认为,当生物-环境系统受到干扰或崩溃时,主导过程将显现。拥有较强环境反馈的系统,将易于快速过渡到新的状态,而由适应主导的过程将改变得较为平缓。反对者同意生物通过生物调节作用影响环境条件以使自身受益,但是生物首先要适应环境条件通过自然选择才能得以繁荣发展的。地球形成这样的环境条件,很可能纯粹是一种运气。(3)自然选择的盖娅:生物调节反馈产生于达尔文式的自然选择。反对者认为,"自然选择支持促进生命效应"的说法并非普遍有效,只有当遗传特征赋予携带者繁殖优势时,自然选择才会支持它。自然选择是机制,而非原则。支持者认为自然选择并不是盖娅系统环境调节的必要条件;基于副产品的自然选择,可以解决许多进化论学者提出的物种合作中的欺骗问题;自然选择并不总是支持促进生命的效应,但在当遗传特征使携带者相对非携带者受益时,自然选择可以使特征携带者产生进化优势。虽然争议依然存在并将持续下去,但作为假说生产者,盖娅假说已经证明了它的价值。但是在人类活动对生物圈影响不断增强的背景下,盖娅假说必须与人类活动相结合,否则必然走向衰落,并被其他理论或假说所替代。在此基础上,未来盖娅假说的研究者们需要继续努力探索可以应用于生物圈的一般性原则,并坚持系统性的思考方法。在具体的方法方面,可以利用系统度量指标;建立新的模型,尤其是建立关于生物地球化学循环过程的机理模型;搞清楚不同尺度过程的成本与收益。     

Early Earth closely surrounded by young stars

Knowledge of the physical and chemical conditions on the primeval Earth is important for the study of the origin of the biosphere. This paper discusses the latest modification of the theory of the origin of the Earth and other planets. Possible consequences of the formation of the Sun in the area of the star formation closely surrounded by neighboring young stars are considered. The classical problem of the rate of accretion of Earth and other planets is generalized with new estimates allowing the correlation of the results from long-lived (U-Pb) and short-lived (Hf-W) space-chronometers. A model of the early evolution of the Earth, based on both dynamic estimates and the latest geochemical data (earliest Australian zircons, relict xenon pleiad) is discussed. The problems of the theory of early Earth’s evolution, which so far cannot be adequately solved, are discussed.     

Kinetic studies on the association and dissociation of myosin subfragment 1 and actin.

The reactions of pyrene-labeled actin with myosin subfragment 1 (S1) and S1-ligand complexes at low ionic strength are described by the schemes [formula: see text] where M refers to a myosin head; A is actin; L is ligand; the asterisk refers to a high fluorescence state of actin; and K1 and K3 are association constants. K1 is reduced approximately 10-fold for M.ADP or M.pyrophosphate versus M alone. The rate constant of the isomerization step (k2) is 150-200 s-1 for A*M, A*M.ADP, and A*M-pyrophosphate (20 degrees C). The interaction between the ligand the actin binding sites reduces K2 from 2,000 for A*M to 50-100 for A*M.ADP and to approximately unity for A*M-pyrophosphate. The A*M.ADP state is equated with the AM'.ADP state of Sleep and Hutton (Sleep, J., A., and Hutton, R. L. (1980) Biochemistry 19, 1276-1283).     

Biogenic Enhancement of Weathering and the Stability of the Ecosphere

A dynamic model of the global carbon cycle is used to determine the influence of biotic amplification of weathering on the overall stability of the biosphere. It takes into account the most important processes for the long-term evolution of the Earth. The model is solved under the condition of slow changing luminosity, volcanic activity, and continental area. We find that for large enough amplification factors the system has two stable states, the abiotic and biotic solution. Furthermore, this leads to an extension of the life span of the biosphere by 0.7 Gyr compared to previous studies underestimating the effect of biogenic enhancement of weathering. It can be shown that the biosphere is resilient to random perturbation of the global carbon cycle for the next 0.5-1.0 Gyr.     

On the early evolutionary stage of the geosphere and biosphere and the problem of early glaciations

The early evolutionary stages of the geosphere and biosphere are determined by three interrelated factors: (1) continuous cooling of the surface and interior (mantle) of the Earth (the mean temperatures of the mantle and surface decreased by a factor of 1.5–2 and 3–4, respectively; the mean heat flow was reduced by approximately one order of magnitude, and viscosity, by three orders); (2) continuous stepwise oxidation of the surface, which was particularly well pronounced from 3.8 to 1.8 Ga; and (3) periodic and correlated fluctuations of conditions in the geosphere and biosphere of varying extent and nature. The major boundaries of this evolution were about 4 Ga (the origin of rather thick and heterogeneous earth’s crust, the origin of life); about 3 Ga (appearance of a strong magnetic field, an increase in photosynthetic activity); about 1.8–1.9 Ga (appearance of an oxidized atmosphere, the first supercontinent, possibly, the first superplumes from the nucleus); and about 0.75 Ga (acceleration of subduction, “watering” of the upper mantle, elevation of continents with vast land masses, shelves, large rivers, and the first great glaciations). The significance and correlations of the earliest events (before and about 4 Ga) and events about 750 Ma are widely debated. In the Late Archean and Early Proterozoic (before 1.8 Ga), the biosphere was dominated by cyanobacteria, the dynamics and developmental peaks of which are marked by the presence of widespread stromatolite buildups in carbonaceous rocks (initially, mostly dolomitic matter). About 700–750 Ma, intense and frequent glaciations developed, marking the cooling of the Earth. The greatest glaciation apparently occurred about 640 Ma, which gave rise to the discussion of the model of the Snowball Earth. The emergence and evolution of skeletons in animals is sometimes thought to be connected with glaciations. These events are correlated and accounted for by great endogenous changes. One of the major events in endogenous history is the onset about 750 Ma of periodic manifestation of mantle flows (superplumes), which explain further periodicity of the biosphere evolution. In conclusion, extrapolation of future evolution and successive collapse of biosphere segments in the course of transformation of the Sun into a red star and warming of the Earth surface are proposed.     

Microbial diversity of deep-sea extremophiles--Piezophiles, Hyperthermophiles, and subsurface microorganisms]

Knowledge of our Planet's biosphere has increased tremendously during the last 10 to 20 years. In the field of Microbiology in particular, scientists have discovered novel "extremophiles", microorganisms capable of living in extreme environments such as highly acidic or alkaline conditions, at high salt concentration, with no oxygen, extreme temperatures (as low as -20 degrees C and as high as 300 degrees C), at high concentrations of heavy metals and in high pressure environments such as the deep-sea. It is apparent that microorganisms can exist in any extreme environment of the Earth, yet already scientists have started to look for life on other planets; the so-called "Exobiology" project. But as yet we have little knowledge of the deep-sea and subsurface biosphere of our own planet. We believe that we should elucidate the Biodiversity of Earth more thoroughly before exploring life on other planets, and these attempts would provide deeper insight into clarifying the existence of extraterrestrial life. We focused on two deep-sea extremophiles in this article; one is "Piezophiles", and another is "Hyperthermophiles". Piezophiles are typical microorganisms adapted to high-pressure and cold temperature environments, and located in deep-sea bottom. Otherwise, hyperthermophiles are living in high temperature environment, and located at around the hydrothermal vent systems in deep-sea. They are not typical deep-sea microorganisms, but they can grow well at high-pressure condition, just like piezophiles. Deming and Baross mentioned that most of the hyperthermophilic archaea isolated from deep-sea hydrothermal vents are able to grow under conditions of high temperature and pressure, and in most cases their optimal pressure for growth was greater than the environmental pressure they were isolated from. It is possible that originally their native environment may have been deeper than the sea floor and that there had to be a deeper biosphere. This implication suggests that the deep-sea hydrothermal vents are the windows to a deep subsurface biosphere. A vast array of chemoautotrophic deep-sea animal communities have been found to exist in cold seep environments, and most of these animals are common with those found in hydrothermal vent environments. Thus, it is possible to consider that the cold seeps are also one of slit windows to a deep subsurface biosphere. We conclude that the deep-sea extremophiles are very closely related into the unseen majority in subsurface biosphere, and the subsurface biosphere probably concerns to consider the "exobiology".     

Two positions in evaluating the role of biodiversity

The necessity is considered to use a compositional approach to the assessment of life phenomena in biosphere instead of, and in practice, in addition to, the traditional reductional approach. From this viewpoint, the construction of logics from top to bottom, from life as an independent system in the biosphere, to system elements, as it is the case in biogeochemistry and the general biological theory, will also appear to be necessary for biogeophysics, if such interdisciplinary science is destined to emerge.     

Origin of biosphere

Concepts of origin of life on the planet are briefly considered. The problem of origin of biosphere is discussed, with a suggestion that the origin of living organisms and biosphere are two aspects of the same process. There is put forward a hypothesis of embryosphere—the primary medium, in which preorganisms could appear. The ecosystemic approach to origin of life poses question about sources of the matter and energy used by the primary life as well as about causes of the biochemical unity that exists in all Earth organisms.     

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.     

Galactic Factors,the Young Sun,the Earth,and the Biophysics of Living Systems

The effects of radiation from the young Sun and galactic cosmic rays on the physical conditions on the early Earth are significantly underestimated in studies of the problems related to the origin and evolution of the biosphere. This review considers the dynamics of solar and galactic processes over the 4.56 billion years of the existence of the Solar System. These factors substantially affected the development of adaptive technologies in ancient and modern living systems. The features of biosphere development are considered for the early Earth under the young Sun, which was fainter, but more flare active. The radiation spectrum of the young Sun is discussed together with the paradoxical mismatch between the solar radiation spectrum and the chlorophyll adsorption spectrum. Ways of solving the paradox are proposed. The role of solar radiation is important when studying models of the early biosphere of the Earth and hypothetical biospheres of giant planet satellites and exoplanets.