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Microbial carpets or algal carpets (Ludwig & Theobald, 1852)

Dans le document OCÉANOGRAPHIQUE Cyanobacteria (Page 193-200)

Geophysiology of cyanobacterial biofilms and the "dyssymmetry" principle

2. Microbial carpets or algal carpets (Ludwig & Theobald, 1852)

3. Stromatolites (i.e. lithified microbial carpets derived from the Greek word stromatactis, the art of making carpets) or stromatolithic mat (Kalkowsky, 1908).

4.Aigai mats, algal skins or algal carpets (Black, 1933, Baas Becking, 1934, Cloud, 1962, Bathurst, 1971, Krumbein & Cohen, 1974; Walter, 1976).

5.Bacterial films or biofilms (ZoBell, 1943).

6.Bacterial and or microbial mats (Doemel & Brock, 1974; Krumbein &

Cohen, 1974).

The term biofilm got into wide use when attachment ofbacteria was studied in caries research and fermenter technology and spread also to natural environ-ments. Ironically the word film does not derive from Greek or Latin but from the Teutonic languages. The term mat appeared as the topic of many volumes and symposia when it became clear that the parahistological skin of the Earth was and is at the basis of life on Earth including rock formation, rock destruc-tion and even plate tectonics and climate shifts. We may thus summarise the impact of cyanobacteria on global processes in the following way:

1. Cyanobacteria are at the base of ail paleontological research of the Pre-cambrian. This allows us to say that cyanobacteria have dominated life on Earth and the shape and dynamics of ail chemical and physical processes on Earth (geological, terrestrial, aquatic, atmospheric) for more than 3000 mil-lions of years.In the last 700 million of years, when eukaryotes spread more

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and more, they doubtless have been the most important factor but now associ-ated with a myriad of interesting emerging macro-organisms which appar-ently aIl are derived from and deeply interconnected with cyanobacteria and their activities.

2.Cyanobacteria are at the base and are also today the major controlling factors of the all-embracing cycles of the elements phosphorus, carbon, oxy-gen, hydrooxy-gen, nitrogen and sulphur in the major geological and global geo-physiological processes.

3.Cyanobacteria contribute to the composition of the gases of the atmos-phere and hereby control the temperature of the Earth (green-house effects, oceanic currents, winds and climate).

4.Cyanobacteria control the productivity of oceans and terrestrial environ-ments to a large extent.

5. Cyanobacteria and other biofilm or microbial mat forming micro-organ-isms control and regulate the composition of the crust of the Earth by creating and depositing carbonate rocks, oxidic (goethite and hematite) and partially or fully reduced (pyrite, magnetite) iron ores and deposits of other metals (metal sulphides) as weIl as siliceous rock types.

6.This way cyanobacteria and associated organisms (living matter) control and regulate the energy content and fluxes of the upper 80 km of the Earth.

7.Cyanobacteria have been and are major factors in the erosion of rocks and mountains (Krumbein, 1969).

The cyanobacterial omnipotence around the globe from the deep sea up to the highest mountain ranges and from the tropics to the poles represents a geophysiological and biogeomorphogenetic power which allows Earth to deviate drastically from theoretical astrophysical planetary equilibria in its atmosphere, hydrosphere, and lithosphere.

Mayer (1845) realised, that organisms regulate and influence thermody-namics to an astonishingly high extent on a global scale. He predicted the modem definition of life, namely the process of drawing negentropy from the environment: "Nature submitted itself to the task to catch the light flooding towards the Earth in its f1ight and to store this most evasive of the forces by binding it into an immotile form. Inorder to achieve this, nature has covered Earth with a carpet" (microbial mat) "of living organisms, which while alive absorb the sunlight and produce a continuous sum of chemical differences under usage of this force."

Only much later it became clear (Vemadsky, 1926, 1928, 1930, 1965;

Krumbein & Schellnhuber, 1990, 1992) that cyanobacteriaIly trapped and immobilised sunlight may even be at the basis of plate tectonics and thus tie together geophysiology and biogeomorphogenesis (Anderson, 1984; Krum-bein, 1990, 1993, 1996). As a conclusion to this sketch of cyanobacterial geo-physiological power we are again attracted by the enigmatic statement of Kant, who opened the debate on "global change" (quoted by Krumbein, 1993): "The mountains of the Earth are sorne of the most useful elements of il. Bumet, who regarded them as a useless and detrimental devastation made 182 Bulletin de l'Institut océanographique. Monaco,naspécial19(1999)

GEOPHYSIOLOGY OF CYANOBACTERIAL BIOFILMS AND THE "DYSSYMMETRY" PRINCIPLE

by God for the punishment of humankind undoubtedly is wrong. According to the method of physicotheology we will describe aIl useful and positive ele-ments of mountains. At the end it might be difficult to decide whether the spherical shape of Earth or the deviations from it belong to the most useful natural facts." Kant also thought that Earthquakes are not divine punishment of humankind but the wise organisation of biogeochemical cycles in order to keep life alive. Herder (1774) went even further and suggested a connection between the living force and the environment in the form of a constant evolu-tionary dialogue: 'The fluctuating and changing climate" (and/or environ-ment) "consists of a multitude of causes which are unequal and thus act slowly and differently until they penetrate into the interior of the organisms and change the latter by adaptation and mutation. The genome withstands this pressure of the environment for a long time, strong in its speciation and self-replicatory power; since the genome, however, is not totally independent from environmental influences, it must be altered necessarily with time."

Admittedly sorne of the terms of Herder, such as "exterior passions" have been transformed into modem language. This transformation was, however, keeping the thought of Herder exactly in the form he has formulated it (Krumbein, 1996). In the second part of this article we will now deal with the history and theory of the dyssymmetry principle.

THE DYSSYMMETRY PRINCIPLE

Pasteur discovered dyssymmetry in the products of wine fermentation he received from a friend and pharmacist. He defined it as "molecular Dyssym-metry". Organic compounds, which are typical for aIl kinds of living matter differ from compounds composing the inert (non-living) parts of the Earth.

He called these two categories: "la nature vivante" and "la nature morte"

(still-life). There are always two enantiomorphs, which could theoretically exist. In the protoplasm of living matter one finds pure steric compounds. In the stereochemical equations of these compounds the atoms preferentially arrange in left-handed or right-handed isomers instead of a statistical distribu-tion as necessitated by physical/chemical laws. Pasteur stated that the bio-chemical processes of living matter and their crystallisation products demonstrate the preferential synthesis and maintenance of left-tuming or right-tuming isomers. He called this phenomenon "Dissymétrie" and defined it as the demarcation line between living and inorganic natural products (Pas-teur, 1922, p. 343). Towards the end of the 19th century the notion of Dys-symmetry was taken up by J.F. Japp (1848-1925) and P. Curie (1859-1906).

Talking about the "directive force" of life Japp (1898, p.453) posed a funda-mental question "whether the phenomena of life are wholly explicable in terms of chemistry and physics" and "whether they are reducible to the prob-lems of the kinetics of atoms". He raised a problem, which is discussed now as the problem of "reductionism" (Rosenberg, 1985, pp. 69-120). In fact, Pas-teur already mentioned "a helical" or "spiral staircase" arrangement of mac-romolecules. Japp proposed that "living matter is constantly performing a certain geometrical feat, which dead matter (inert matter) is incapable of per-forming". He formulated a general principle in his "Stereochemistry and

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Vitalism": "Only asymmetry can beget asymmetry." In view ofthis confusion between asymmetry of Japp and "dissymétrie" of Pasteur we will now use the term dyssymmetry. The prefix "dis" may mean "different or other than", but also "double" as in "bi" or "bis". Symmetry originally cornes from the Greek language and thus we prefer the Greek prefix "dys" with its meaning of separate, different, other than. We regard symmetry and asymmetry as a pair to which dyssymmetry is added as deviating from but not being the opposite.

This agrees with the mathematical/morphogenetic approach ofD'Arcy Thomp-son (first published 1917, 1984 edition usually used) to organisms and living matter.

Japp (1898, p. 458) said: "We thus arrive at the conclusion that the produc-tion of single asymmetric compounds, or their isolaproduc-tion from the mixture of their enantiomorphs, is, as Pasteur firmly held, the prerogative of life. Only the living organism with its asymmetric tissues, or the asymmetric products of the living organism, or the living intelligence with its conception of asym-metry, can produce this result. Only asymmetry can beget asymmetry." Japp (1898, p. 459) also asked: "How could léevo-rotatory proteins be spontane-ously generated in the relatively symmetric non-living world, if we accept that the principle of symmetry is one of the fundamentals of nature?"Ifby accident left or right handed enantiomers were produced in excess in a purely chemical reaction then they wouId equilibrate by racemization. This pheno-menon by itself is still another blow against spontaneous generation (Krum-bein, 1995).Ifdyssymmetry is not restricted to life then a hitherto unexplored principle of the "nature morte" should be responsible for the sudden or grad-uai tum to dyssymmetry in living matter. Therefore we can postulate once more that life or the "Iife field" is a principle of nature (Krumbein, 1983a, Cramer, 1993).

P. Curie used the original terminology of Pasteur. He wrote that, if sorne effects manifest a dyssymmetry, then the same dyssymmetry must be observ-able in the causes of these effects, although the opposite is wrong, i.e. the results can be more symmetric than the causes of this results (Curie, 1908).

Curie connected this principle with the idea of a "state of space" (état de l'espace) and coined this term as a potentially different view of the living and non-living world.

Vemadsky reports about a conversation with M. Curie: "She thinks that it is this notion (état de l'espace) that contains the synthesis of ail his thought (Vemadsky, 1965, p. 160). M. Curie was convinced, thatP.Curie just did not have enough time to elaborate this idea, because of his sudden death in 1906.

" The notion of "state of space" defines space as a real existing entity. State of space is determined by physicallchemical or biophysicallchemical events.

P. Curie realised that, if space is something that really exists, then the causes and their effects must find themselves in the same space. Applied to the prob-lem of dyssymmetry this means, that if phenomena and processes exist in space, where left and right are not identical, then the causes could exist only in the dyssymmetrical state of space. This ties the problem of dyssymmetry closely to the question of abiogenesis (spontaneous generation or "omne vivo e vivo").

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Vernadsky created a mature theory of dyssymmetry and elaborated a theory of spatial-temporal dyssymmetry of living matter. Inthe following we discuss only Vernadsky's concept of spatial dyssymmetry. This concept is a part of his theory of biological space and implies two meanings of the term dyssymmetry: (1) a phenomenon called dyssymmetry of life (Urmantsev, 1992, p. 25), of which the dyssymmetry of protoplasm (Pasteur's dyssymme-try) is a special case and (2) dyssymmetry of the biosphere.

On pure mathematical grounds Vemadsky considered those objects as dys-symmetric, which:

a. are not coinciding with their mirror images (geometrically, chemically, energetically);

b. can exist exclusively or preferentially against thermodynamic considera-tions in either of the two enantiomorphic modificaconsidera-tions (left and right);

c. exist in one or two different modifications selectively and preferentially expressed in biological systems or living matter.

The dyssymmetry of Pasteur is the peculiar case of the molecular dyssym-metry of life. Vemadsky talks of the dyssymdyssym-metry of Pasteur, when sorne chemical compounds typical for a living natural body statistically can and should exist in two modifications but exist only in one modification during the whole natural history of life (Vemadsky, 1965, p. 198). When the second modification is artificially synthesised it manifests the difference from the natural modification by its properties. Evidently, the latter statement is trivial and does not allow us to talk about the dyssymmetric character of space

"occupied" by living matter.Inorder to substantiate this idea Vemadsky takes up the notion of the "state of space" of P. Curie in addition to Pasteur's origi-nal articles. Vemadsky proposed a principle, which he called the Curie princi-pie: "Dissymmetrical effects (phenomena) can be brought about only by a dissymmetrical cause." He realised that, if space is a kind of intelligible real-ity, then the causes and their effects must find themselves in the same state of space, i.e. they must be embraced by a certain state of space (Vemadsky, 1965, p.182).Itwas remarked later, that the addition made by Vemadsky is so important that the latter principle should be called the Curie-Vemadsky prin-ciple (Eliseev, 1989, p. 196).

Different states of space can be more or less separated, but also close to each other (Vemadsky, 1965, p. 169). Symmetry is a criterium of a state of space: "Symmetry characterises the different states of the Earth space" (Ver-nadsky, 1965, p. 169). The "state of space" of any natural body can be deter-mined by the basic principle of symmetry. This principle declares that the state of space of a natural body is determined by the minimum symmetry of its properties (Vemadsky, 1988, p. 379).

Thus the dyssymmetry of Pasteur corresponds to a special case of symme-try breaking, becauseitis completely out of the tradition al laws of symmetry of the non-living natural world. The dyssymmetry of living matter can be seen as a very special case of asymmetry and/or symmetry. Using the term

"state of space", the "basic principle of symmetry", and the characteristics of dyssymmetry we can perceive that the "state of space of a living natural

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body" will be characterised as dyssymmetric, although sorne elements of its structure can be symmetric or asymmetric in a non-dyssymmetric way.

Vemadsky emphasised, that his approach to space and space-time differs from the physical-mathematical approach ofA. Einstein. The "space" of the relativity theory is a "space of measurements", whereas the space ofVemadsky-Curie is a "space of states". The "space of life" or as a whole the "life field" is characterised by the dyssymmetrical state of space, where the leftness and rightness and other factors are not symmetrical (identical) or statistically dis-tributed (Vemadsky, 1997, pp. 217-218). In living matter the properties of "to be Jeff' and "to be right" are not identical. The dissymmetric state of space of living matter is of a hereditary nature and reproduces itself during millions of years. The destruction or annihilation of this state of space means death.Itis one of the reasons for the necessity of multiplication: The dyssymmetrical state of space cannot be achieved by the non-dyssymmetrical states of space.

The problem of the dyssymmetry of life implies not only the chemical dys-symmetry of protoplasm (dysdys-symmetry of Pasteur).Itfurther implies the dys-symmetry of isotopes and other atoms, which are selectively collected by living natural bodies. Age dating of organic matter as weil as the attribute

"biogenic" to certain types of ore deposits and other natural phenomena are related to these novel types of dyssymmetry introduced by Vernadsky. The preferential selection of certain isotopes and elements by biological systems is only possible under the terms of extracting negentropy from celestial sun energy. Galimov (1989) states, that Vemadsky was the first to realize, that dyssymmetry applies also to these essential factors of living systems. He wrote: "Actually, the ferments organise the space in which the movement of biological matter takes place in a special way." Therefore it is possible to assume, that the ideas and the notions ofVemadsky have been realised (albeit in slightly different terms and categories) in the contemporary theory of the biological fractionation of isotopes (Galimov, 1989, p. 343). Vernadsky fur-ther includes the question of the relative abundance of sorne biologically important enantiomorphs in the discussion. He expanded the Pasteur-Curie principJe of dyssymmetry into new atomistic (isotopic) and geometric direc-tions. The degree of morphological dyssymmetry can further be connected with the level of entropy (Velikbekov, 1969, p. 14). Vemadsky realised, that the chemical dyssymmetry of Pasteur had to be distinguished from the mor-phological dyssymmetry. He further proposed a connection between these two kinds of dyssymmetry. Experimental work in this line has been initiated by Gause (1940) He studied left- and right-spiralled colonies of Bacillus mycoides,which were compared with the structures of enzymes.

Vemadsky's "morphological" and Pasteur's "chemical" (protoplasmic) dyssymmetry embrace the phenomenon of dyssymmetry of life. Dyssymme-try of life is deeply connected to the problem of the peculiar character of the biological space. Biological space is physically different from the inert/non-living space. There seems to be a mathematical paradoxon between the inert/non-living and non-living world, which cannot be solved on the basis of quantum chem-istry (Bohr-SchrOdinger-Delbrück-paradoxon). The physical space-time sce-nario is reversible but it seems that the biological space-time scenerio is irreversibJe as characterised by the evolution or emergence phenomenon.

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The principle of symmetry is refracted not only by the prism of dyssym-metry, but also by the characteristics of symmetry in living matter. The number of symmetry elements observable in any solid (crystal, mineraI) of the inert matter is restricted. In the living world one can observe bioobjects with forbidden for the crystals axes of symmetry of 5,7,8,9 order. The min-eralogist W. Bragg pointed out that there are no regular dodecahedrons among the natural crystals.Incontrast to this it is possible to show symmetry axes of the sixth and seventh order in living organisms (Vemadsky, 1988, p. 264; 1965, p. 190). The up-to-date level of the life sciences makes it possi-ble to describe samples with even much higher structural symmetry than Ver-nadsky mentioned. Contemporary investigations of morphological symmetry show that we can find in living matter not only dodecahedrons (Circorherma dodecahedra) but also macrobio-objects with symmetry axes of 5, 7, 8, 9, etc., orders, which are not realised in the mineraI world. Urmantsev (1974) stated, that the macro-level of biosymmetry also demonstrates the correctness ofVemadsky's thesis of a peculiar character of the biologieal space. Urmant-sev generalised a vast set of empirical data concluding that in the case of living matter both on the macro-Ievel (evolutionary) and on the micro-Ievel (cellular) distinct dyssymetrisation can be observed (Urmantsev, 1974, pp. 211,218). Vernadsky, however, mentioned two main kinds of dyssymme-try, which we c1assify as: dyssymmetry of life and (morphogenetic) dyssym-metry of the biosphere.

Dyssymmetry of life describes one of the main properties of living matter in the theory of Vernadsky, which implies chemical dyssymmetry of Pasteur and morphological dyssymmetry (micro- and macro- levels in the classifica-tion of Urmantsev, 1974). The space of a living natural body is determined by the presence of dyssymmetrical elements. We are stating dyssymmetry when there is an element in the structure of a natural body, which exists only in a preferred enantiomorphic form; another enantiomorph, being statistically demanded for or even theoretically admissible, must show different essential properties. Vernadsky further postulated, that dyssymmetry can exist also out-side of the immediate field of life of a single organism. It will be, so to say, a dyssymmetry of the second order,because it can be seen as a consequence of the activity of ail living matter on Earth (the biosphere).

This "secondary" dyssymmetry is observable in the structure of biogenic substances (e.g. organic fossils and organic matter as petroleum, coal and evaporative hydrocarbons). This includes the question of how long a biogenic product such as petroleum would keep dyssymmetry elements after being released from the body of a living being (specimen, population). Racemiza-tion of amino acids e.g. is a time/space related physical/chernical phenome-non of slow reactions of radicals. This racemization reaction, however will only start and get significant when the compound in question has been released from a given living natural body (ceIl, organism, biosphere).

On the global scale we find another phenomenon: Dyssymmetry of the geosphere: "Not ail geological envelopes and geospheres are ideally round", although there are no pure geological reasons for this fact (Vernadsky, 1965, p.llO). Considering the form of our geoid (or better bioid), we can expect ail geospheres to be ideally round with the deviations permissible by planetary

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