7. MUTUAL INFLUENCE

“Nothing in the world is single” P.A. Shelley.

There ought to be some debate on the question whether the mutual influence of organisms constitutes a section in Geobiology. It certainly has nothing directly to do with the direct influence of organisms on the earth’s crust. Still mutual influence as will follow from the consideration in this section, is for a larger part chemical influence and there is, therefore, a continuous exchange of substances, not only between inanimate milieu and the organisms, but also between different organisms. This exchange of substances might even, in some cases, directly influence the outward environment. Even when it doesn’t apparently influence the milieu this might be due to lacunae in our observations or to inimitable material. The great influence of symbiosis upon geobiology is the shape of the biosphere, the form it takes, its natural composition, as the components of a symbiosis and of a biocoenosis form a sort of a symphony orchestra. Organisms are linked up together, as in a gigantic tissue the pattern the warp and the woof. We cannot segregate one actor from this play, one voice from this chorus, without disturbing the ensemble and therefore, when continuing biology here, it should be synbiology.

[In the margin Baas Becking referred to Section 6.3, Periodic Changes in Chemical Milieu; Section 7.2.1, Table 7.2a and 7.2b, Symbiosis; Fig. 7.14 showing interactions between microorganisms and higher animals.]

Fundamental to any independent existence of a living unit is the power of carbohydrate synthesis, a synthesis which pre-supposes the faculty of carbon dioxide and oxygen. This faculty, as we have seen, is given to the green cell and, moreover to various groups of colourless bacteria. While the green cells use the sunlight as a source of energy, the colourless autotrophs utilise external energy. Carbohydrate synthesis is fundamental to the independent being, but to all living beings should be common the power of plasmic synthesis, at least to a certain degree. We may consider the glucose sugar as a starting point and some here in its metabolites, e.g., the ketoacid³ consider an amination (according to Martius and Knoop (1937) and Martius (1937) first to organic acid, then to imino acid and finally to amino acid).⁴

These are the first steps of plasmic synthesis. About further stages we know even less. It may be said however that, according to Bergmann, every proteinaceous molecule is built on a recurrent pattern of certain amino acids.⁵ The ideas of B.C.P. Jansen, according to whom the pattern is developed on a certain matrix, and, in themselves not only probable, but obvious.⁶ The glucose metabolite, as well as the energy necessary for the basic and functional maintenance of life are derived from the glucose molecule. In order that the vital functions are performed normally, this sugar metabolism should not be interfered with. We have therefore, as these fundamental metabolic processes:

1. The anabolism of glucose (A),

2. The catabolism of glucose (K),

3. The plasmic synthesis (P).

A green plant cell is mostly AKP, a higher animal akP or aKP. Now it is obvious that the latter organisms are unable to persist without an organism AKP or AK. The organism aKP is dependent upon cells which are able to synthesise sugar. It has been customary, however, since the days of De Bary, to designate as symbiosis another relation between organisms, which might be symbolised as Ab (organism 1) and aB (organism 2) living together. Each of the components, in themselves, are unable to persist independently – together however they form a “higher unit” AB, which is capable of existence. It is a case of “the lame that leads the blind.”

It seems, however, arbitrarily, to narrowly delimit the concept of symbiosis and to restrict it to cases of mutual benefit. For at the basis of symbiosis (in the strict sense) and nearly all other interrelations of cells and organisms, there is the process of material, or chemical exchange. We shall have occasion to consider seemingly distantly related processes from under the high viewpoint of symbiosis. It may be seen that in many cases even the rigid definition of De Bary seems, after closer inspection not applicable. The “simple” food relation grass → cow for instance, may well be considered as a classical example of symbiosis. For, in the country of Holland, where forest is climax, meadows are artificial landscapes, caused by grazing animals. [inserted: Barbed wire causes the grassland to become a dense alder thicket within a few years.]⁷ The grazing animal itself, therefore creates the conditions necessary for the development of his chief food. There would be no grassland without cattle, there would be no cattle without grassland.

Still, the old definition does not pertain to such phenomena as parasitism and to the antagonistic relations between organisms. Neither does it hold for organisms that change the milieu in such a way that others may follow a succedaneous⁸ rather than a simultaneous symbiosis (metabiosis, term suggested to me by H.G. Derx).⁹ But it seems to express a general rule in the higher animal organism in which, withincreasing morphological differentiation, the chemical power of individual tissues or cells seem to decrease, causing a complicated condition of mutual dependence. Before we attempt to classify the various phenomena of interdependence, it may be well to describe, anecdotally, certain cases of symbiosis. And we will return to our first example: the cow and the grass. The green cells of the grass blade¹⁰ are able to perform all sorts of inorganic reductions: CO₂→glucose, hexavalent S→bivalent S, nitrate to ammonia. The cells of the grass blade, moreover, are able to break down the formed sugar without external aid, such as aneurin. At least, if we consider such species of grass which are free from mycorrhiza. Furthermore, the cells of the grass blade are able to synthesise, out of the glucose catabolites and ammonia, all amino acids necessary for the formation of the protein molecule.

Most of the photosynthate is stored, however, as cellulose, which forms by far the greatest part of the hay. The cow, at the other hand, is unable to produce sugar from carbon dioxide. It is equally unable to produce glucose from cellulose. It cannot, without external help, metabolise sugar. It needs the phytonic principles, the vitamins.¹¹ Without the carotene, which is part of its visual purple, it cannot see. And finally, it is only able to synthesise a few amino acids, while most of these “protein building blocks” have to be provided for. With the great morphological differentiation which has led to the organism we call “cow”, there has been an increasing chemical importance. A cow + grass is, therefore not even a “closed system” in the sense of Beauverie and Monchal (1932). We need the cellular bacteria and the protozoan population of the stomach in order to decompose celluloses and, perhaps to form the necessary amino acids, enzymes, ergones (vitamins) necessary for the maintenance and the growth of the cow. The inability of the cow to decompose cellulose may be considered as a deficiency in glucose metabolism. The cow, the grass and the microbial flora and fauna of the bovine stomach form a higher, symbiotic entity. The components of such an entity influence our milieu. Nearly every organism is tied to others by many bonds. Nothing in the world is single. [As a note Baas Becking added: It should be specially stated that evolution cannot be conceived as performed by individual species. Due to the interdependence of organisms, evolution should be a process in which genetically unrelated but symbiontically related species, should vary simultaneously. Tentatively this concept is named synevolution.]¹²

A second example is taken from quite another field of biology (Fig. 7.2). Let circle b in the accompanying figure represent the milieu possibilities of a terrestrial orchid, let us say an Ophrys. Quite dependent upon this Ophrys is a rust, the milieu of which is depicted by circle a. A digger wasp, showing a high degree of mimicry (Føyn),¹³ performs cross pollination. Its milieu boundary c is about identical to that of the orchid. Finally, we know that the endotropic mycorrhiza, the Corticium living in the subterranaean parts of the orchid, is not specific, but may enter into symbiontic relations with other plants. Therefore, circle d, denoting the milieu boundary of this Corticium, exceeds the others. Here the complex is formed by four components, it is still one of the very single examples of symbiosis.

The deficiency of the functional performance of an organism has to be completed and overcome by another organism or its products. The three large functional groups mentioned: assimilation, dissimilation and plasmic synthesis, the chemical nature of which is obvious, has to be completed by other groups, to wit, sexuality and development. Here also we meet with chemical effectors (Geschlechtsstoffe, Hartmann and Moewus;¹⁴ organisers, Spemann;¹⁵ Needham).¹⁶ The complex of these substances we shall designate by S and D respectively. The group S, however, is in certain cases, not a “conditio sine qua non” for the existence of an organism.

AKPSD is the most complete, and the most completely independent organism, akpsd the most dependent. As said before, the higher animals belong to the latter groups, where concomitant with morphological differentiation and speciation, chemical performance has materially deteriorated. The higher animals need besides its trophon, its caloric food, a great number of ergones. It is quite probable that for the five functional groups mentioned, a large part of the substances needed are synthesised, and provided for, by plants. So, for groups K the B and C vitamins are necessary, for S the vitamin E, while for D the D groups are of paramount importance. Now if we consider a single cell possessing the properties AKPS and D.

This cell gives rise, by division, to various groups of tissues, deficient in one or more of these functions. The organism, while still being as a higher unit, capable of all these functions, represents a consortium of specialised cells. Such a consortium we will call an autosymbiosis. If the consortium is built up of various species, we may speak of heterosymbiosis.¹⁷ Gamosymbiosis is the sexual relation between individuals of the same kind, and therefore a form of autosymbiosis. Sociosymbiosis, the formation of a socium contains both auto- and heterosymbiotic relations. Saprophytism and parasitism also fulfil the requirements as belong to the wider concept of symbiosis. A parasite autosymbiosis is the formation of the embryo within the mother organism. Saprophytism, in most cases belongs to the metabiosis or succedaneous symbiosis. In order to oversee the difficulty of a “contradiction in terminis” the word coenobiosis may be used to cover both the simultaneous (symbiosis) as well as the succedaneous (metabiosis).

The expression of coenobiosis also termed biocoenosis, a community of various organisms which are more or less in mutual equilibrium with each other and with their internal milieu. It stands to reason that the concept of symbiosis lies at the basis of the knowledge of a biocoenosis and that the chemical and physical process as well. A biocoenosis leave their imprint upon the biosphere. The knowledge of the mutual relations between organisms is therefore of paramount importance to the geobiologist. At the end of this section various biocoenosis shall be, as far as possible, analysed.

We may inquire a little further into the potencies of organisms. A simplified scheme is given below (Fig. 7.3a). Carbon dioxide assimilation [anabolism] is represented over an intermediary A. Dissimulation [catabolism] over an intermediary C. Cellulose formation and breakdown also represented. Further reduction of nitrate and sulphate, formation of amino acid and formation of protein.

Scheme Coenobiosis:

* “penicillin” and “oxaline”,¹⁸ modern natural bacteroides and probably nothing but “defense substances” of fungi. Such defence substances we also find in the seeds of Capparis and Tropaeolum and within the glands of Humulus fruits (all bacterial).

A complete autotroph may be represented by Figure 7.3b, that is the abstracted scheme from Figure 7.3a.

Now Quispel (1943 and 1946) proved that the lichen Xanthoria has gonidia which cannot perform photosynthesis without a diënol. This alga is therefore (1) or (2). The fungus needs aneurine, is therefore (4). They are therefore as follows (roman numerals mean “capable”, arabic numerals is “incapable”):

The cow, the grass and the bacteria:

See for this case Section 7.4.4, Phylosymbiosis.

The organic counterpart of the minimum substance is the “Wirkstoff”, the ergone, the nutrilite, or by what other names we want to call an ever increasing host of organic substances, each of them claiming to be necessary for ‘something.’ It is beautiful to work with an ergone, as one obtains specific effect from a specific substance. Pharmacology does this too and it has been, therefore, happy with it. But if biology be reduced to a host of little bottles full of ergones each of them ‘doing something’, while we have not the faintest inkling of the “why”, there is a great danger in the modern pharmacological riot which seems to reign in biology.

Let us recall the examination at the end of Le Malade Imaginaire of Molière. Here the board asks why opium puts a fellow to sleep: the considerate answer that there is a specific virtue in it, “Bene, bene, respondere.”²² We feel as helpless as the modern medic who applies the new patent residues with their beautiful names. Apart from the dienoles and vitamin A, I have not the faintest idea how an ergone or enzyme or serum, or hormone, works. It works and we know that is the most important. For it may be that, what we call “neutral influence” is directly the work of ergones and that this entire living nature is linked together by them like a gigantic organism. The question may even be put, whether there is an intrinsic difference between ergone of the enzyme, hormones, vitamins and the like.

When we know more about the specific function of ergones, the ‘pattern’ of each symbiosis may be written down. So, the classical case of Schöpfer and Jung (1937) see 7.5.19. Unfortunately, chemical analysis has not proceeded far enough to allow us to carry the matter further. H.G. Derx, 15-8-44 in litteris.²³

The excrements of all kinds of animals are particularly rich in ergones. Haematococcus pluviatilis grows best on cow manure. We think about the work of Thaxter on Myxobacteriaceae,²⁴ which he found on all sorts of dung (rat, rabbit, etc.) and several genera of fungi, named in Saccardo (in finis bovis etc.) (strumania).²⁵ It is interesting to think that the organic substances, after passing through the intestinal tract should not only be assimilated but that, in their stead, microbial as well as “host” ergones should appear in the excrement. Asteromonasgracilis Atari and Brachiomonas both Polyblepharids, occur in salt pools with bird excrement. Thinking of the work of Darwin on the earth worm one wonders what this animal contributes in the way of ergones! Bassalik (1913) isolated the bacterium Pseudomonas extorquence from its gut, a bacterium able to decompose oxalic acid.²⁶ ‘Coprology’ may be a promising science indeed! (See for this Section 3.12.3a, Humus). Suggested experimentation with excrements of a great variety of animals, sterile filtrates and tried in pure cultures of a number of standard organisms.

Vitamin A, reaction Carr and Price (1926) (SbCl₃).²⁷ ½ molecule of 𝛾 carotene.

[Baas Becking inserted Fig. 7.4.]

Provitamin A, Vitamin A, anti-endophthalmitis²⁸ (carotene part of the visual purple).

Goldberger ‘black tongue’, ‘pellagra’ factor.²⁹

[Baas Becking inserted Fig. 7.5.]

Aneurin, Co carboxylase, thiasole.

E. Wildiers found already in 1901 that few yeast cells hardly served as inoculum, but many cells gave good result (controversy Liebig/Pasteur).³¹

Ascorbic acid C₆H₆O₈,

Reductive acid (von Euler),

dioxymalic acid,

reduction,

are all characterised both possession of the dienole group,

which may be oxidised,

all these substances have an abnormally low redox potential. Ascorbic acid was discovered by Szent-Gyögyi, it is now industrially made from sorbate.³² Dioxymaleic acid is more out tartanic acid with H₂O₂. In glucose solutions sterilised at pH > 8.0 sometimes appears which has an unstable Pb salt. Reductive acid was also discovered by v. Euler in pectin substances.³³ They all react with di-ether phenols indophenole, but very slowly, with methylene blue (after illumination). Ascorbic acid is the ascorbic vitamin. The ascorbic acid water as a mediator in oxidation. Dioxymaleic acid, moreover, seems to destroy oxydases.

Ascorbic acid is present in most green plants. Very much in paprika, green peas, strawberries, asparagus and cauliflower stem. Giroud claims that it accumulates in the green plastids but the reaction used by him (AgNO₃ in light) is hardly to be called specific.³⁴ However, Quispel (1943) proved that certain lichen gonidia could only perform photosynthesis when ascorbic (or for that matter) dioxymaleic acid were present (in a hydrogen atmosphere). It may be that in plants the dienoles function as oxidation regulators and, moreover, as a kind of photosynthesis ‘hormone’.³⁵

Vitamin D1 and D7 anthraquinone derivates. Closely related toad poison, Arginea [Micropsalliota arginea] and Digitalis glycoside (Stoll, 1937),³⁶ sex hormones (Butenandt, Ružička) and carcinogenic substances.³⁷ It is claimed that the animal sterols are synthesised by the animal cell (cysterol vs. phytosterole). This seems improbable.

See Nicolai in Vakbl. v. Biologen (1943).

Indolyt acetic acid and butynic acid, the heteroauxines = rhizopon.³⁸ See Koningsberger, Leerboek derAlgemeene Plantkunde (1943).

Since Went’s classical work in 1927,³⁹ and the brilliant chemical researches of Kögl, we know of plant hormones that are able to make cellulose walls plastic in that they may be extended by longer pressure until new molecules of cellulose be laid in between the old (internode sections). Higher plants produce the auxins proper, microbes and bluegreens also heteroauxine. Auxine may be inactivated by light (lumiauxin).⁴⁰

Vitamin E contains, apart from the chinons a long phytol chain, identical to that met with in the chlorophyll molecule. Evans proved the presence of the fertility vitamin in wheat germ 1929.⁴¹

See also 5.11.1, Anaerobiosis, Aerobiosis; 6.4.2.a, The Composition of the Atmosphere.

Vitamin F (Burr and Burr, 1929),⁴² and a skin vitamin, the absence of which in the food causes “plagues” to appear in rats. It has been shown that the substance is probably linoleic or linoleic acid. A rather large quantity is needed (identity with fearon acid!).⁴³

See Figure 1.1, Figure 7.14 and Scheme Coenobiosis in Section 7.1.

Let us consider a higher plant in its symbiotic relations (see Tables 7.2a and 7.2b).

• Of ktenosis we only find phagoktenosis (4) in insectivorous plants.

• Oiko-ktenosis here (6) maybe at the basis of the substances like penicillin or oxaline and the germination preventing substances of Fröschel. For these, and allied matters see Funke (1942).⁴⁵

• Histo-parasitism we find in the embryo (7). Gamo-parasitism is typical for all higher plants, as the sexual cells both few on surrounding tissue, especially the pollen tube. (8).

• Ecto- and endoparasites (10,11) we find galore, wasps and plant lice may cause galls, to them a case of oikoparasitism (13). In the differentiated higher plant, there is histo-helotism (e.g.,  host tissue) (14), while the fly trap flower, like Aristolochia, actually exploit insects (gamo-helotism 19).

• Seeds with spines etc., promote the oiko-helotism (20) by which animals disperse the seed without water! Histo-mutualism describes further the chemical correlation on the plant body (21), while the sexual act is seen as gamo-mutualism (22).

• The growing together of plants of the same species may be an expression of ortho-mutualism (23).

• Ecto- (24) and endomutualism comprises the classical cases of symbiosis, while the so called flower biology is named here as gamo-mutualism.

• Seed dispersal when the seed is eatable or eatable in part (ornitho-chorisis, myrmica-chorisis),⁴⁶ may be called oiko-mutualism (27).

• Plants may, on dying give of useful substances (28) to the species (necro-symbiosis 28), they may live a saprophytic life (ecto-saprobiosis 29), while their ontogeny (32) and the succession of species inwards and external (33) are cases of metabiosis.

• In the complex of symbiosis plants already represent more than twenty cases of sapro-biosis!

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Increase in specialisation when traveling upwards in the evolutionary scale, from bacteria via algae and mosses to ferns and further to higher plants, we find an increasing degree of specialisation of the individual cell, capable to perform all functions.

The cell in Figure 7.6 shows a cross section of a root of a higher plant and a section through the stoma of a leaf, to show differentiation.

Table 7.3 shows, approximately the number of cell types while ascending the evolution ladder in the plant kingdom. For the animals a similar series could be built up. Now it is impossible to take a cell of a root parenchyma and raise a whole plant out of it. Only the epidermal cell of e.g., the Begonia has kept its “prospective potency” (Hartsema, 1926) and may generate a whole plant. Most of the cells are too far specialised to perform this feat. The entire plant body is able to perform that what the green unicellular did by itself. Morphological differentiation has decreased the chemical potentiality, leaving the total potency the same or even increased. From this it follows that autosymbiosis should be considered as essentially biochemical in nature, and that development and differentiation go hand in hand with continuous and synchronous exchange of ergones.⁴⁸

J. Needham, chemical embryology, Spemann (organic concept).⁴⁹

Most tissue cultures of animal cells, revert, in the long run, to connective tissue, Differentiation ‘goes back’, retrogrades; there is de-differentiation and differentiation (Gaillard).⁵⁰ Tissue cultures from chicken embryos have shown that differentiation of, let us say bone tissue, only occurs when juice has been added from a chicken embryo of the age in which it forms bone. This shows that differentiation is induced by certain very specific substances. The same holds for thyroid, parathyroid.⁵¹ This the first direct proof of the existence of ‘organisers’ throughout the entire developmental cycle. It seems not too far fetched to claim that throughout the life of organisms, in removal and in breakdown of tissue, such like substances should be continually active.

Endocrines, when synthesised by the organism itself, should be considered in the same way as the mycetomes (Buchner, 1921) of insects.⁵² Here we have islands of tissue, the cells of which are microbes or filled with microbes, which cells secrete substances necessary for the maintenance of the organism and which substances are transported by body fluids (‘𝜊𝜌𝜌𝛼𝜔’ I carry a message).⁵³ It is a fanciful, but not entirely creative, thought to consider the endocrine glands of higher animals as evoluated mycetomes. It is of course known, how all the messengers carry ergones, as secretin, adrenaline, choline, thyroxin, progestron, androsterone, pituitrine etc. Again, here there is quite a mass of ergones! And this increase in the number of these endo-ergones in evolution goes hand in hand with morphological differentiations. The more highly differentiated, the more dependent the cell becomes upon its environment.

1) Isogamones (Polypodiophyta).

2) Mobile heterogamones (brown and green algae).

3) Sperm and egg (animals, insects, Nereis, ferns).

4) Pollen tube and Ambyocea (higher plants).

5) Conjugation in Conjugales (Spirogyra).

6) Conjugation in Phycomycetes.

7) Diplo-haploid [= diploid] mycelium in Eumycetes.

The scheme in Figure 7.7 gives a few types of gamosymbiosis, which end, in any case, with the formation of a zygote. In case 4, two coalescence products of nuclei are formed the zygote + the endosperm molecule. The latter being triploid. In Zygomycetes the + and – myceline grow towards one another attracted by substances secreted by the partner. It is probable that also the pollen tubes are led by a chemical stimulus. The school of M. Hartmann in Berlin, has made a profound study of sexual behaviour in lower organisms, believes in the chemical basis of mutual attraction of gamones.

The chemotaxis of the moss and of the Pteridophytes sperm was first discovered by Pfeffer (1884) and Stahl (1884).⁵⁴ Here again, different chemical substances (sugars in the case of mosses), malic acid in Equisetum, citrine acid in ferns, were the substances secreted by the cells near the ovarium in order to attract the sperms. Apart from the chemical substances which play a part in the secondary sexual characteristic of organisms (obvious) we must assume, in higher animals, the presence of chemotactical substances secreted by the female.

Moewus, Wendt and Kuhn (1938), have found that, in Chlamodomonas eugametos, not only sexual behaviour but also mobility was brought about by traces of substances generated by the algae, derivates of crocine, a carotenoid, the cis-and trans-crocetine corresponding to “attraction” substances for male and female respectively. ⁵⁵ The old experiments of J. Loeb should also be recalled in this connection.⁵⁶ Sexual behaviour of cells, gamosymbiosis, is shown to have a chemical basis and differs in no way from other forms of symbiosis.

There are two important questions before us: primo, is symbiosis possible without evolution and, secundo, the inverse, is evolution possible without symbiosis? The second question was raised by Noel Bernard (1899) later by Bernard and Magrou (1911) and answered in the affirmative.⁵⁸ It shall be seen that both questions are closely linked, and that it appears as inconceivable that organisms evoluated by themselves. There should be a “syn-symbiosis”: a cow is really ‘a grass consuming engine.’ Its jaws, its dentation, its stomach, its gut, it is a grass mill. There is a symbiosis grass ⇆ cow, rather one sided, but very real. There is also a symbiosis (as a sort of afterthought) bacteria ⇆ cow, cellulose bacteria, without which the cow would not digest the grass. She maybe, needs the bacteria also to supply her with ergones, or complete protein, or both. In the triangle cow ⇆ grass ⇆ bacteria there is only one autotroph: the grass. There would be no cow without grass. The cow presupposes grass. The tapeworm presupposes a root, the orchid presupposes a digger wasp, the ladybird comes after the aphid, the koala after the Eucalypt.

Evolution therefore, has to be syn-evolution. And while the thought of this type of evolution first matured in the brain of a (hereditarily Lamarckistic) Frenchman, we claim that Lamarckism has never had its proper experimental chance. It is one thing, like Weismann did to cut off mice’s tails and try to raise tailless offspring, or to paint salamanders with India ink, like Kammerer, or to consider the multitude of ergones, keeping in mind that we already know of naturally occurring substances, like colchicine, that may change the genome of the offspring. It cannot be stipulated emphatically enough that evolution is an anabasis, not of a single specimen, but of a closed mass of various organisms. And that it well may be that changes in the genome, which lie at the base of evolution, are brought about by ergone action. That the excitant, as is known living in symbiosis with the variant, had to look for other fields of endeavour. This is not a neo-Lamarckian that looks to the inanimate environment as an evolutionary agent, but it is a Lamarckian that claims that the Evolution of organisms, is caused by organisms apart from the fact that we need creation. The agent of which, according to Spinoza is as active now as ever before, with which beautiful thought the author heartily concurs.⁵⁹

In Figure 7.8 the scutellum is covered by a glandular epithelium known in germination to secrete amylase⁶⁰ However, it may secrete a great number of things important for the young embryo that has for a considerable period to be fed from the endosperm.

The sporogon is entirely parasitic upon the gamotophyte in liverworts (Campbell, 1905),⁶¹ and in mosses Although the sporogon is arranged with a conductive tissue and with stomates it sucks, by means of a foot, all of its nutrition from the gametophyte. Its CO₂ assimilation, however, is unimpaired. In the lycopod the prothallus may be subterraneous and saprophytic. Here the diploid plant is actually and completely parasitic for years, before it reaches the light (Büchner, 1897a and b). In the forms the case is, in the beginning, somewhat like the mosses. Even in a highly developed plant, as the pine, the young embryo is fed by the primary endosperm, which is nothing but a prothallus, part of the gamotophyte.

In the flowering, so called higher plant, before fertilisation we get the reduction divisions of the spore mother cells and the formation of pollen and of the embryo sac. The remnants of the haploid prothallium, which is entirely dependent upon the surrounding tissue of the (diploid) sporophyte. This pertains already in the heterospore Lycopodiaceae (Selagonella) and persist throughout Cycads and Gymnospenia although in these forms the gametophyte is still highly developed. We do not know what type of exchange there exist between haploid and host.

(Seeds in plants, usual embryology). In higher plants the young embryo is either surrounded by the diploid cotyledons, like in the legumes, or one find an endospore, which consists of triploid cells. This is, in a way, an organism itself. We find here reserve food, oil, or starch or proteinaceous. On germination this reserve food is mobilised. An egg (bird) is, in a way also a diploid embryo capable of living apart. There is a very complicated parasite relation between embryo and mother, in which exchange of ergones is really the crux of the whole problem of embryology.

(Ezekiel 27 Faraoh).⁶²

Politeia is more than a biocoenosis, it is a number of organisms in such a close community of intent [intentional community] that the obvious analogy with the human state suggests the name given above. Of course, a politeia occurs only where social animals are domiciliated for an appreciable time. There is a great difference between a colony and a politeia. In a colony (e.g., reef coral) community of intent (cilias, movement to move the water) unite, but the components are not actively engaged in promoting it. Ants, bees and wasps and termites are the communities we are thinking about in the first place. There is no doubt that among the ants the civitas is most highly developed. Amongst the higher animals we know nothing of the kind. Schools, flocks, coveys, packs, herds are all concepts which convey a quite different meaning. Only in Homo sapiens there reappears politeia, not as a natural thing, but as an afterthought, perhaps as a necessity. The prospective potency of the components is, in human society at least, unimpaired (of Begonia leaf, Hartsema, 1926).⁶³ In animal politeia the smith would have hammer instead of hands, and the carpenter perhaps a saw or chisel. Our plasticity may be, however, not externally fixed.

In a way, the syn-evolution proposed by Bernard (1899) and by Bernard and Magrou (1911), is nothing novel. It is apparent that the organisms in themselves form a ‘web of life’ just as much as they, naturally form such a tissue. We are driven, nolens volens, to a novel form of Lamarckism, an evolution induced by ergones. We see, in the ascending series, an increasing amount of chemical dependence, of ‘chemical helplessness’, going hand in hand with an increase in morphological and functional differentiation. However, in the plant kingdom, there is another type of evolution. Here the organism, while developing morphologically, has kept its prospective potencies.

The two-toed sloth Bradypus has hairs with a furrow, in which a green filamentous alga lives (see Weber-van Bosse, 1887).⁷¹ Mammals should not be green, like lizards or caterpillars, for they are able to metabolise the chlorophyll entirely. Therefore, when my friend Walter Spies told me that he had shot a green bat on the island Nusa Perida, south of Bali,⁷² I expected a symbiotic alga. As a specimen of the bat was present at the Buitenzorg collection (Dobsonia viridis), v. Bemmel,⁷³ assistant curator examined the hairs and actually found a bluegreen alga within its hollows, bifurcated hair. The observation was extended to Dobsonia hair obtained from the Amsterdam Zoological Museum, through the kindness of its curator L.F. de Beaufort.⁷⁴ Here also the alga was found. This is a case of an epiphytic alga, an arbitrary relation between two organisms. Now the relations between organisms are manifold (see Section 5.2.3). In the following pages we shall deal chiefly with mutualistic symbiosis. A full description of the cases would require an extraordinary amount of space. Mutual influence is so pronounced that we have used the term ‘web of life’ to it (see Section 1.4, Fig. 1.1, Section 4.9.5 and Tables 7.2a and 7.2b).

ArdisiaMiehe (1911 and 1917), de Jongh (1938), Bok.⁷⁵ [See also Section 6.4.2.b]

[Baas Becking inserted Fig. 7.9.]

Mej J. van Roon.⁷⁶

Rayner.⁷⁷

Hellriegel and Willfahrt (1888), Beijerinck (1888), Fred, Allen, Löhnis (1941), Virtanen.⁷⁸

[Baas Becking inserted Fig. 7.10.]

Roberg (1934 and 1938), v. Oven, Honbold [?].⁸⁰

[Baas Becking inserted Fig. 7.11.]

Reinke (1873), Hansen, Baas Becking (1947a) and Hulssen (1946).⁸¹

In all of the Cycadaceae inspected by me in the Botanic Garden Buitenzorg Java 1939, as with Cycas, Encephalartos, Zamia, Macrozamia, Stangeria, Dioon,⁸² the roots, especially those near the surface, show nodular excrescences which, on closer examination show dark green under the surface (Fig. 7.12). Bluegreen algae, intracellular are the cause of these nodules and of the colour. Sometimes the colour is non-existent, the cells seem therefore to contain still another organism apart from Nostoccycadae. Bacteria have been repeatedly found within the cells. Experimentation is still lacking, but it may be that, like in the case of Gunnera, root formation is stimulated by the bluegreens (+bacteria?) according to Galestin (1933),⁸³Nostoccycadae to capable of assimilation of atmospheric nitrogen. Obviously, when this were the case, it may be that, apart from ergones, the Cycas is supplied with amino acids, which for some reason it needs.

[Baas Becking inserted Fig. 7.13.]

De Bary (1864 and 1879), Schneider (1898), Jacq, Quispel (1943 and 1946).⁸⁴

Orchids.

Bernard (1909), Rumpff (1741-1750).⁸⁵

Treub, Jacq., Campbell (1905), Bruchmann (1906).⁸⁶

Azolla.

Straszburger. ⁸⁷

[Baas Becking left this section blank.]

Corals, sea anemones, hydra’s, sponges, de Laubenfels (1930), Boschma, Verwey.⁸⁸

In unicellulars, protozoa such as Paramoecium bursaria⁸⁹ and Stentor amethystinus⁹⁰ we meet with unicellular green algae, which one named (by their negative characteristics) Chlorella. In other protozoa, foraminifera and radiolaria, we meet with yellow symbiotic algae, Chrysidella. In higher forms such as hydra’s, anemone’s, sponges, corals we always speak about Zooxantellae, and the colours of the symbiotic algae have may be, as a matter of fact, either green or orange-brown. The latter colour is caused by the phenomenon of climatisation and is a typical deficiency phenomenon.⁹¹ Evidently, in this case, the host has not been able to meet the requirements of the symbiont (⁠$NO3−$⁠, $H2PO4−$⁠). A great many algae, (Polyblepharids, Protococcales, Conjugates) when brought in adverse conditions, will develop their carotenoid pigments in anomalous quantities (see Vreede, 1941?).⁹²

Buchner (1921) in his classical treatise has shown that a great many insects contain certain glandular structures in or near the gut which contain bacteria, yeasts or other fungi.⁹⁴ It is surprising that blood drinking insects are often provided with mycetomes, showing that, apparently, they do not derive all of the necessary ergones from the host. Apart from the mycetomes, they apparently have other sources of ergones. The aphids, for instance, reach a sexual period often when the host plant starts flowering. If this is a short day plant, the aphid shows two periods annually (Aphis forbesi on strawberry; MacKoritch cited in Shelford, 1929).⁹⁵

The Hypermastigniae, an order of the flagellate protozoa, are symbiotic with termites and with woodroaches such as Cryptocercus (Kudo, Fig. 119-131). There are a great many of these protozoa for a large part described by Kofoid and Swezy.⁹⁶ The adult termite does not attack wood; it lives on the excrements of the larvae. The larva is able to destroy wood or rather, as Cleveland has demonstrated,⁹⁷ the fauna of the gut, whether solely protozoa in character or in combination with bacteria, is able to digest cellulose. Here we meet with a close analogue with what we find in ruminants.

The Australian marsupial, our well known Koala, lives on the leaves of very special Eucalypts. In a ruminant and compound stomach the leaves undergo an alcoholic fermentation, banning the animal to be more or less continuously sleepy and, unfortunately cross. Here symbiosis should probably include a yeast, although bacteria may also generate alcohol from sugar.⁹⁸

Galls.

W. Schöpfer was the first, in 1934, to realise an artificial symbiosis between a red yeast Torularubra and a phycomycete Phycomyces blakesleeanus. Both organisms are heterotrophic and, moreover, aneurine deficient. Torula is able to synthesise the thiazole half; Phycomyces the pyridoxine half of the molecule. They are therefore, only together a stable combination. This finding opened our eyes to the real significance of symbiosis and not much later Kögl and Fries (1937), see also Funke (1943).⁹⁹

Walking leaf, branch, Mantis (there is a crustacea like this).

Bug Reduvius.

For merfly like bumblebee.¹⁰⁰

Mottled background fishes, salamander.

Velvet ants.¹⁰¹

Orchids and wasps.

Moths.

Birds eggs.

Winter pelts in mammals.

Geometer caterpillars.

[Baas Becking inserted Fig. 7.14.]

Here we meet with a succession of organisms, the medium begins presequent by the sequent for the subsequent.

As there seems to be a certain contradiction in the term succedaneous symbiosis. The author shall follow a suggestion of H.G. Derx and use the term metabiosis.¹⁰⁴ The question remains whether there exists true metabiosis. Of course, all actors are present, only behind scenes or, biologically speaking, in latent form. According to some, and the number of examples is countless, they are always present also in the vegetative stage, but their numbers are small (examples may be given from lactic acid bacteria, etc.). There are instances, however, of true metabiosis, where the milieu has to be prepared and for organism B by organism A.

Intestinal symbionts are known to synthesise several ergones. The rat, for example, does not need vitamin C, it prepares it in its gut, or rather its microfauna prepares the ascorbic acid. (Giroud, L’acide Ascorbique).¹⁰⁵ Now it is remarkable that many microbes live on manure. It is said that the former Professor of Botany at Amsterdam, C.A.J.A. Oudemans, already in the ‘sixties’ gave a course in microbiology.¹⁰⁶ To this end horse manure was put under a bell jar, and the developing flora observed daily! (See also Lindner, 1888, Gärunggewerben).¹⁰⁷ First Mucor appears, then Thamnidium elegans, followed by Pilobolus, Sordaria firmicola, while the series ends with Coprinus. Now this may very well be a true metabiosis, but on a very rich nutrient medium, in which many ergones are already present! Cow manure seems to be particularly rich in ergones, as the work of H.G. Derx (in litteris) shows. Thaxter has shown that Myxobacteria are typical for rat and rabbit dung, while Humaria, Peziza and other Discomycetes are common on cow and deer excrements.¹⁰⁸ Saprophytes on humus, also a complicated, ergone-rich, mixture, is, of course better known. It is possible that a weird practice of the anthroposophist’s, called “biological-dynamical” method of manuring, in which a compost is prepared with many ingredients, really is efficient because of the number of ergones it introduces into a metabiosis. The formation of the so called “Edelkompost” should therefore remain a biological process and never be degraded to a chemical manipulation, as several modern patents claim to make “humus” out of refuse at high temperatures with NH₃ etc.

Already in 1908 Pütter claimed that there should be a certain amount of soluble organic matter in natural water which might serve as food. The Wisconsin survey has investigated this matter and found indeed in Wisconsin lakes organic material in hue solutions as the following Table 7.4 shows.¹¹⁰ However we know now, especially after the work of Moewus and Wendt that organic substances functioning as ergones are active in very low concentrations (---γ/L for excretion) so that Pütter’s hypothesis obtains renewed significance, not so much as a source of caloric food, but as a source of nutrilites.¹¹¹ This necro-symbiosis when hetero-symbiotic, may be succedaneous.

The literature on the brine bacteria claims that there are freshwater forms living at low temperatures. As I found empty sheaths of these bacteria at solar salt water near Bombay and also on the island of Madura, this claim seemed erroneous. As I was unable to culture the Gallionella directly (except in freshwater at low temperatures), I used the methods of succedaneous culture, which is an example of metabiosis. Use was made of filter paper pulp and in organic medium, infection was with diluted mould, the whole jar kept anaerobical. At first there appeared cellulose bacteria, anaerobic forms, which at first made some gas but soon ceased in doing so when the next group, the sulphate reduction bacteria appeared. Due to the presence of ferro-iron, ferro-sulphide (FeS) or troilite was formed until the contents of the entire jar became pit black. Now air was admitted and then within a few days the aerobic sulphur bacteria appeared, which changed the S, formed by the non-biological oxidation of FeS, to sulphate. When moreover the bottle was placed in the light, development of purple bacteria could be seen within 10 days at low temperature. Now the conditions for development of iron bacteria were fulfilled! There was Fe²⁺ and oxygen, at the surface of the synthetic mould they appeared, at 35 °C, in strong brine!

Now this method suggests a great number of possibilities and a number of microbes that have been refractory to cultivation might develop in the media (Harmsen).¹¹² Apparently the succedaneous (or cyclic) culture originates subsequently a number of ergones which number increases as the culture proceeds. It would be a beautiful field of research to study such succedaneous phenomena. Cellulose fermentation is a promising starting point, as in nature cellulose materials often function as “initiators” of cycles.

(Söhngen and pupils).¹¹³

(Bacteria followed by phage). After the discovery by Twort and Hérelle of the bacteriophage,¹¹⁴ agricultural bacteriologists have demonstrated that lytic agents occur in soil bacteria as well. Radiobacter beyerinckii, nitrifies and other bacteria show the typical “parasite” in culture. It has been assumed that protracted sterility of soils, such as are met in the Residency of Indramaju in Java, might be caused by the development of certain soil bacteria. The concept is somewhat too much simplified as soil fertility implies an enormous biotic cycle in which both metabiotic and symbiotic phenomena should occur.

Biological purification, whether in rivers or in septic tanks is a typical metabiosis. Spoilage, of course, is a more or less anthropocentric term, but it may mean an accumulation of organic substances in water, changing thereby its character in such way that it becomes unfit as potable water, industrial or fish water. The accumulation of organic matter, however, incites various swaps of microbes to great activity and mineralisationetc.

The classic case is the purification of the Illinois River below Chicago (Kofoid, 1903),¹¹⁵ or the septic tank and filter system. Here first enormous numbers of bacteria develop, which, in their turn, are consummated by protozoa. If the water is sufficiently aerated, the end products of mineralisation (protein, fats, sugar) are nitrates, sulphate and carbon dioxide. Even so called antiseptics, like phenol, are finally broken down. There are fungi however, certain cases, where mineralisation is incomplete (sulphite lignin from paper factories). Bacteriophage also may play a great role (Ganges, Nile). When taking in new drinking water aboard a ship (Wibaut-Isebree Moens, 1916) first bacteria develop.¹¹⁶ Within a week they are cleared away by protozoa, leaving the water clear and harmless to health.

(Backer, 1929; Docters v. Leeuwen, 1936; Funke, 1943).¹¹⁷

In the so called Krakatoa problem, we meet with the initial vegetation on sterile lava. In our laboratory a method was conceived by Baas Becking, later by de Jongh and elaborated by Quispel (1938).¹¹⁸ A “Zaponlac” film is prepared on top of (not too humid) soil and this film fixed by means of “Geisselthal-lack”, after thorough drying the film is removed. (The method originated from the chance observation. While preparing plaster casts of the tracks of game animals, the author found a film of algae adhering to the plaster, which could be satisfactorily microscopied). The surface vegetation may now be studied in situ. Even clean white quartz boulders yielded fine preparations of moss protonema and Zygogonium ericetorum.¹¹⁹ By means of this method Quispel found that, on sterilised lava at temperatures of 30 °C first diatoms and then moss protonema appears. The bluegreens come much later. It would be an excellent field of investigation to study succession ab initio. (To study the surface film of water, the same lack is used. A brass ring, suspended by 3 strings, is dipped into the lack and then held at the surface of the water) (see Fig. 7.15).

A sterile lava is really the only habitat where we may speak about succession of organisms, as in other cases we always meet with simultaneous elements as well. This is also the case with plant succession, which are dealt with at length in the sociological works of Blytt-Sernander and of Braun-Blanquet. It is the hypothesis of Funke (1943) that these successions are just another expression of production and exchange of certain symbiotic substances, or the stimulation of certain bacteria, which may prepare the soil for the next phase. ¹²⁰ In van Dieren (1934, Organogenic Dune Formation),¹²¹ we first get the beach plants, adapted to moving sand, then the plants of the outer dunes, rich in lime sand, as the CaCO₃ is leached out, the plants of the lime-free, inner dunes, which culminate in a heather and the heather may yield to oak-birch forest. It seems logical that a concomitant change in the soil microbes should take place.

[Baas Becking inserted Fig. 7.16. On top of the Figure: “Besemer.”]¹²³

A parasite is chemically very helpless. In certain textbooks it is claimed that it cannot e.g., synthesise its proteins, and therefore is so dependent upon its host. Now proteins are specific and as parent and host usually are phylogenetically totally unrelated it seems far fetched to claim any affinity between their proteins. Most probably the parasite has to break down the host’s protein entirely and then start to build its own. For this breakdown as well as for this building it may miss chemical tools. It may miss the tools to break down sugar too. Therefore, a reinvestigation of the problem of parasitism by biochemical means seems promising.

[Baas Becking left this section blank.]

A most remarkable characteristic of many parasites is their often far going dedifferentiation, which goes hand in hand with inability at chemical synthesis, which we consider to be the basis of parasitisation. The rule that morphological differentiation occurs at the price of chemical dedifferentiation has to be amended somewhat. The higher plant is morphologically and chemically highly differentiated (MC), the lower plant cell would be mC, the higher animal Mc, while the parasite should have a symbol mc. The dedifferentiation we meet in the parasites in the ticks (Ixodes), worms (Taenia), crustaceans (Sacculina), and many insects (e.g.,  swallow fly). In plants this dedifferentiation is not so marked, although Santalum, Cuscuta, e.g., show morphological regression. Dedifferentiation does not always accompany parasitism, however, not is it restricted to parasitism, as saprophytes (Latraea) and even epiphytes (Taeniophyllum) show morphological regression. It seems that in both cases the host cannot apply the variety of enzymes necessary for normal functioning.

Sprirochaetes. Spirochaeta pallida cannot be cultured on artificial media, on fresh blood substrates it is possible. What ergones are the cause of the phenomenon is not known. The live entirely in dissolved substances, as there is no mouth. Amoebae are capable of phagocytising; they live chiefly on bacteria. Myxamoebae have been cultured by Schure (Thesis, Leiden, 1935).¹²⁴Entamoeba coli, Councilmania lafleuri, the latter cause of amoebial dysentery. The question remains whether they only accompany (and few upon) pathogenic bacteria. Ciliates. A great many ciliates are always present in the intestinal tract of animals. Pathogenicity is often indicated but extremely hard to prove, as the organism cannot be cultured. Certain free living ciliates have been cultured by Woodruff (1914 and 1921), Flagellates, Lamblia internalis, Giardia in man.¹²⁵

Leeches are provided with mycetomes. Many blood sucking organisms have mycetomes. Blood seems to be insufficient diet to them. Leeches secrete hirudin, a substance prohibitory to coagulation of the blood of a host.

Roundworms. Ancylostoma duodenale, mice worm,¹²⁶Oxyuris vermicularis,¹²⁷ in the sphincter anus region, Ascaris lumbricoides, roundworm, in many mammals. Capable of an oxybiosis, may be cultured outside the host. The dedifferentiation is not marked, structure as complicated as in free living Nematodes.

Tapeworms. Taenia, Bothriocephalus, have been kept alive in protein solutions of unknown composition.

Lice contain mycetomes. Pediculus vestimenti (typhus vector) [Baas Becking added a note: had I only known! 31-V-45]

Phthirus inguinalis,¹²⁸Pediculus capitis.¹²⁹

Flea Pulex irritans (plague vector Yersinia), mycetomes. ¹³⁰

Bugs. Cimex lenticularis, contains mycetomes. Bugs are probably vectors of many bacterial and protozoal diseases.

Tabanids.¹³¹

Mosquitos, Stegomyia, Anopheles.

Nearly all pathogenic bacteria may be cultured on artificial media. However, they are not strictly defined media, as one is made of blood cells.

Tuberculosis B. tuberculosis, [modern name Mycobacteriumtuberculosis] many characteristics of actinomycetes.

Diphtheria. B. loeffleri,¹³² many characteristics of actinomycetes.

Plague. Yersinapestis, semi treatment recently successful (Otten, 1936).¹³³

Cholera. Vibrio cholerae. Semi treatment successful.

Typhoid. B. typhorus.¹³⁴ Semi treatment successful. See for literature on ergones A. Den Dooren de Jong, Vakblad v. Biol. 1942.¹³⁵

Actinomyces, Madura foot,¹³⁶ may be cultured, although growth is slow (Lüske) [not identified]. Probably highly incompetent in synthesising milieu interne.

Dusts, fire, mildew

Skin fungus is usually cultured on saturated medium (malt extract + peptone Poulenc). Rijkebrinck [not identified] showed that protein may be replaced by ammonium salt. What ergones are needed is unknown.

Cuscuta.¹³⁷

Loranthaceae.¹³⁸

Santalales.¹³⁹

Cycle and Metabiosis. A cycle is a sequence of phenomena which show recurrency. The old emblem of the snake with its tail in its mouth symbolises the cycle aptly. The cycle may be polar, one end representing aquatic life, the other terrestrial, or one side a high energy level and the other a low energy level. There may be components that are (temporary or permanently) removed from its influence (‘slugs’) but its recurrence is not broken by these processes. Cyclic processes suggest a sequence of events (See Cyclic or succedaneous cultures, Section 7.6.4). So, the circle itself may represents time (of day, of year, of month). The circle may also represent energy level, as in the case of the material compounds, centring around a certain element. But still, the time affect persists, as all these processes take time. In some cycles the links are strictly succedaneously in others, all acts are staged simultaneously. In some instances, the first link prepares the way for the second, and so on, until the circle (and the cycle) is closed. Then we meet with a real instance of metabiosis, as defined by H.G. Derx. But in this way the life cycle. In bacterial cycle and even the vegetation cycle may be linked, as in all cases there is dependence upon material exchange of a more or less definite nature.

In this section we shall deal with the so called “successions” of higher organisms first. They are too complex to be analysed as yet, they may only be described. Geochemically they are very important, as they determine the nature of the biosphere cover. We shall, furthermore, consider typical seasonal cycles (annual, monthly, diurnal) as they determine the majority of milieu influenced biological phenomena. Cycles of longer range we shall not mention here. In the third place we shall deal with material cycles. First of certain elements and compounds in their mutual relations, finally about three elements in particular: carbon, nitrogen and sulphur in which centre a great number of important geochemical reactions. Their relation to hydrogen, oxygen and electron transfer will be shown.

We further refer to Section 7.8.8 of this treatise.

(Deli, three fallow system)

[Baas Becking inserted Fig. 7.17.]

As to agriculture and horticulture references should be made of alternation and fallowing and other recurrent procedures.

Further literature should be consulted. (Some of the more sensible plant ecology could be worked in here).

Forest is climax in our latitude. It should be stated which milieu condition, further the formation of grassland.

[Baas Becking inserted Fig. 7.18.]

Diurnal cycles in plants, animals, man.

Annual cycles in plant, animal and man.

Diurnal milieu cycles.

Annual milieu cycles.

Tidal cycles (see therefore Section ---)

Tidal worms from Wilhelmshafen.¹⁴¹

Opening of stomata, purple bacteria, daily and annual rhythm in animal and human activity.

Cambial activity.¹⁴²

This to be elaborated when the pertinent literature has been perused!

“Des Menschen Seele gleicht ganz dem Wasser. Zum Himmel geht es, vom Himmel kommt es, und wieder nieder zur Erde muß es, ewig wechselnd” Goethe.¹⁴³

a. Iodine. Kroprapport Nederland 1921 (iodine).¹⁴⁴

b. Water. See Geobiologie (Baas Becking, 1934).

c. Iron. See Correns (1939). Fe³⁺, Fe²⁺, porphyrium, FeOH²⁺, ferrites, ferrates, and oxides FeO, Fe₂O₃, Fe₃O₄ and hydrated FeS, FeS₂.

d. Oxygen.

Absorbed by weathering of rocks (1)

Oxidation of FeS and FeS₂ (2)

Oxidation of organic matter (3)

Respiration of animals and plants (4)

Oxidation of other inorganic matter (5)

Liberated

By photosynthesis (6)

By chemosynthesis (7)

Balance see Section 5.12.4

Iron, minerals, haematite Fe₂O₃, limonite Fe₂O₃.H₂O, siderite FeCO₃, melanterite FeSO₄.7H₂O, magnetite Fe₃O₄.

[Baas Becking inserted Figs. 7.19 and 7.20.]

To describe the carbon cycle in one page is tantamount to reviewing the largest part of the living, for carbon is the essential element of living beings. The starting point is the atmospheric CO₂, about the notation of which we refer to Figure 7.21 and Figure 7.3. The picture below is self explanatory. The highest reduction to methane, requires the donation of 8 electrons. This is the highest energy level of the cycle. In CO₂ assimilation only 4 electrons are transferred to the carbon dioxide. Humification and caramelisation are probably non-biological phenomena (see Section 3.12.3a, Section Humus). It is the question whether the universally occurring oxalic acid could not be derived from 2 CO + H₂O₂ !

The energy relations as given in Plate 3.3 see Section 3.6.10.

There is only one such a diagram because C^4- and C⁴⁺ which oppositely changed, have the same valency. The existence of a C and a CO organism is questionable.

[Baas Becking further referred to Plate 3.1, see Section 3.5.15 in which the glucose breakdown is graphically represented. He also added Fig. 7.21 to the text.]

Maximally 4 molecules of water may be transferred to 1 CO₂, yielding methane CH₄. In CO₂ assimilation only 2 molecules of water are fixed upon CO₂ with the formation of 1/6 C₆H₁₂O₆. Here the C is half reduced. To the transfer of every two electrons then correspond 1 molecule of water. In order to find the reduction state (n of I electrons transferred on C⁴⁺) we use

This number, multiplied by 28,200, (1/2 ∆H, H₂O) yields the heat of combustion (approx.) of the compound (see Plate 3.2) It is of course improbable to mention here the components of the carbon cycle. They embrace the large part of biochemistry. Only a few specific remarks should suffice.

1) Porter (1926) has claimed to have found an organism oxidising C.¹⁴⁶ Coal on heaps loses 10 % of its weight in one year. This may also be inorganic (pyrophonic iron) However, process seems to stop after application of disinfectants.

2) CO, the presence of CO ergones by no means proven.

3) CH₂=CH₂ may become very important! Ripening of fruit etc.

‘Slugs’ in the cycles.

There remain in the cycles compounds that do not enter the cycle again readily, or only when certain conditions are fulfilled. Some of these products and their conditions are:

[Baas Becking inserted Fig 7.22.]

Clarke (1916, Geochemistry) assumes that all of the organic nitrogen should be derived from the atmosphere. Fixation of the nitrogen starts the cycle. There are, as stated at other places in this treatise, a symbiontic aerobic Radiobacteria, further and independent aerobic Azotobacter and an anaerobic Closteridium, a butyric acid ferment and a few Cyanophyceae, Nostoc? Oscillatorium, all able to reduce the nitrogen, probably all with the aid of molybdenum as a catalyst. The efficiency of the Azotobacter is much less than the Haber Process (Baas Becking and Parks, 1927). Probably hydroxylamine is an intermediary compound. Further we find, as an autotroph Winogradsky’s Nitroso- and Nitrobacteria (Kingma Boltjes, 1934) proved that they are facultative heterotrophs) and denitrifying, producing N₂ and N₂O from NO₃. The green plant and various bacteria are capable to reduce $NO3−$ to NH₃. The NH₃ enters either keto acid, to form, via immuno- and amino oxides, the aminoacids (Martius and Knoop, 1937) or they are accompanied by oxal acetic acid (vitamins) to form asparagin and glutamin (Chibnall, 1939). ¹⁴⁸

In the reduction eight electrons are again added to N⁵⁺. The composition of the biologically important H, N, O compounds are given in the two diagrams, constructed for N^3- and for N⁵⁺ respectively (see Fig. 7.23a and 7.23b). As the energy relations are particularly complex, they are not included in the diagram. The line N-H₂O probably represents the area of lowest potential. In the lower triangle the reactions NH₃ to HNO₂ and HNO₂ to HNO₃ are given as hydrogenation of ammonia, resp. nitrous acid (see Fig. 7.3a and 7.3b). The nitrifiers have an efficiency of ±5 %.

As far as nitrogen fixation is concerned, the diagram for N⁵⁺ gives this in terms of hydration and dehydrogenation of N₂, taking hydroxylamine as an intermediary product. As nitrogen has very little affinity for water, the above equations are a good example of paper chemistry, however.

[In the margin:] The book of Correns (1939, p. 241) recognises only the aerobic part of the nitrogen cycle and the conclusions for N⁵⁺ reaction about the Zechstein are unwarranted.

[Baas Becking inserted Fig. 7.24.]

Geochemically most abundant is the sulphate, we shall start with that compound. It is reduced back by green plants and by anaerobic bacteria to sulphide ion, which on exposure to the air yields free sulphur. But H₂S, aerobically may generate S by autotrophic aerobes.¹⁵⁰ Anaerobically, the reaction is endothermic, and the organisms which perform this feat are photosynthetic (purple and green bacteria). While the reaction H₂S + O = H₂O + S is chemical, the reaction $S→SO42−$⁠, is mainly biochemical. In this reaction aerobic autotrophics may go as far as a thiosulphate, but usually (Thiobacillus thiooxidans)¹⁵¹ they go the whole road to sulphate, of the forming free sulphuric acid. In the green plant we find sulphur as thiocyanate, mercaptum¹⁵² or disulphite sulphur and in aminoacid.

The triangles in Figure 7.25 refer again to S^2- and S⁶⁺ respectively. In the complete reduction 8 electrons are again added. The energy relations are opposite to those in N₂, S being the highest level, as illustrated in the overall triangle in Figure 7.25.

Sulphide oxidation and sulphate reduction may be both represented by hydrations and dehydrogenations or the inverse, as seen from the diagram and from the accompanying set of equations. The excellent paper by Bunker (1936, Ministry of Publ. Health reports of the United Kingdom) should be consulted and the various organisms noted. ¹⁵³ The energy efficiency of sulphur bacteria is again low (5%). It is worthy of note that the sulphate reducer, when grown on a substrate containing only mineral salt and hydrogen, behaves as an autotroph (D. Stephenson,¹⁵⁴ compare the work on the corrosion of iron pipes of von Wolzogen Kühr).¹⁵⁵

Sulphate reduction makes oxygen available, and the oxidation of S → SO₄ is concomitant with the reduction of CO₂! However, as the efficiency of these autotrophs is only 5-10 % the geochemical balance will not be disturbed materially by these processes. Other sulphides, such as those of zinc and copper, may very well originate in a similar fashion as the FeS, and also the oxidation of these sulphides to sulphate may be bacterial (see Section 5.8, Minimum Elements) From all bacteria living in the organic substrates, the sulphur bacteria probably are geochemically the most important. Their action, especially in the ocean, has been described (see Section 6.4.4). Consensus of opinion among geologists exists as to the large sulphur deposits (Sicily, Louisiana) as of being of biological origin. Much sulphate (gypsum) should also be of microbiological origin, despite the dictum of Correns (1939), that all of the gypsum is marine in origin.

Green plants and chemosynthetic bacteria function as the only inorganic reductants, but the quintessence of this function lies in the power to photolyse or to electrolyse water, to reduce the hydrogen oxide! Here the electron has to be fixed into the H⁺. In the case of metallic iron in the soil this process may be understandable (Fe + 2H⁺ → Fe²⁺ + 2H), but when there is no electric element the mechanism of the process remains obscure. It also may be that the electron moves from OH^- to H⁺ according to

It may be that there is still a deeper relation between the heat of formation of water and the energy values of organic compounds (Plate 3.1 and Plate 3.2). It seems that ½ of the energy is necessary to add one electron to a cation. The matter still requires looking in to further (L. Pauling, 1939, The Chemical Bond).¹⁵⁶

Cycles of C, N and S are a series of dove tailoring oxido-reductions, endo- and exothermic, each linked with its organism or groups of organisms, a veritable exposé of metabiosis, of succedaneous symbiosis.

A biocoenosis is a natural community consisting of various vital components. Biocoenosises may be, like organisms, autotrophic or heterotrophic.¹⁵⁷ Whenever they are self supporting (containing a preponderance of autotrophic components), they form a real closed community. Heterotropic biocoenosises may never be understood without taking into consideration their source of energy. The biocoenosis may be, therefore a closed universe, repeating on a small scale the story of the biosphere in its entity. However, milieu conditions are now specialised and the natural milieu is much narrower than that on earth. Our milieu of the biocoenosis should be specific enough to characterise the organisms expected to live here. The simultaneous occurrence, on the other hand, of several specific organisms characterises a specific biocoenosis. By the organisms found in a brine lake one is able to predict (within bounds) the chemical composition of its waters, while, when knowing H and S analysis, one is able to predict the occurrence of its inhabitants.

In 1925 the author enclosed in a quart bottle some natural brine of approx. 20 %, containing the eggs of the crustacean Artemia, the unicellular alga Dunaliella, various protozoa and bacteria. The bottle was ¾ filled and hermetically sealed. The community remained active for over four years, having yielded countless generations of Artemiae. After that time the bottle was filled with a dense sediment, the skins of these Phyllopods consisting of chitin which, apparently could not be decomposed with sufficient velocity. A sample of freshwater from a dune lake, containing Ostracods and bluegreen algae, collected 1937, remained active for 3 years. Here again the chitin accumulated. The study of these “polycomponent” systems still yields results not amenable to analysis, but if two or three component systems were studied, one could, by increasing the complexity step by step, probably arrive at a much better understanding of natural biocoenosis (see Beauverie and Monchal, C.R. 1932, 195).¹⁵⁸ The aquarium is, in a way, a closed universe, as Warrington¹⁵⁹ and especially J. v. Liebig have recognised (Liebig, Die Welt).¹⁶⁰ By a universe we mean a part of the biosphere, where creative and destructive metabolism (catabolism and anabolism), Shiva and Vishnu,¹⁶¹ are both active.

The Amsterdam Zoo Natura Artis Magistra contains several sea water aquaria, constructed in 1900 and in 1933 studied by Catharina Honing.¹⁶² The aquarium is an ideal biocoenosis to study and has been used quite often as an example of natural cycle. The Amsterdam sea water aquaria contain copepods in the water which is derived from the conduits. Due to the oligodynamic action of these copepods the flora is extremely limited. There is, therefore, the fauna and the food, together with a filter system of sand, and an aeration system. Anaerobiosis apparently does not occur (only small patches of black mud observed), but necessary reduction processes are also hindered. Deficient in the cycle is denitrification. There is an accumulation of nitrate, due to the fact that the “salpetre micrococcus” of Beijerinck cannot perform its function, as the oxygen tension is too high, due to continuous aeration.

The community of the black mud, FeS, troilite imbedded in silt, clay or sand is characterised by an extreme anaerobiosis. The community consists of the Sporovibrio desulfuricans,¹⁶³ cellulose bacteria, Clostridium, of green organisms, certain bluegreens (Phormidium) and Euglenids (Eutreptiaviridis), further several protozoa and nematodes. The origin of the black mud is not as Hecht (1933) and Correns (1939) still seem to believe,¹⁶⁴ the decomposition of proteins as Dorothy Stephenson has raised, Sporovibrio as hydrogen and energy source.¹⁶⁵ The author of this essay found most copious sulphate reduction in perfectly inorganic media (Melianthus [?] solution) with finely divided filter paper. Here cellulose fermentation sets in first and the sulphate reduction liberates as the fatty acid produced (Baars, 1930), or as the hydrogen produced. Metallic iron as Wolzogen Kühr showed,¹⁶⁶ forms, with the soil solution an electric element, giving rise to iron ions and free hydrogen. The latter yields the energy needed for the reduction of the $SO42−$ according to

The third reaction being Fe²⁺ + S^2- → FeS

See further Section 2.3.2.

As soon as oxygen is admitted an entirely new biocoenosis appears also when light is admitted. Pyrite FeS₂ is of secondary origin according to FeS + S = FeS₂ (Verhoop, 1940) and is of non-biological origin. The black mud may be one of the largest ‘oxygen traps’ on earth, as during oxidation large quantities of 2FeS + 3O → F₂O₃ + 2S oxygen are fixed. ¹⁶⁷

[In the margin: The community is dependent, in most. It needs hydrogen as fuel and therefore usually, organic food.]

1. Amoebae.

2. Diatoms.

3. Flies.

4. Beetles.

5. Bluegreens.

6. Sulphur bacteria.

• a) Ruppia maritima (Walfish Bay!).¹⁶⁸

• b) Artemia.¹⁶⁹

• c) Dunaliella – Ephydra.

Artificial brine (Boekelo).¹⁷⁰

a) Lochmiopsis – Brachionus.

b) Dunaliella – Ephydra.

Varves.¹⁷¹

Varves.

[See also Geobiologie, 1934, p. 110 English edition, 2016.]

Varves.

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Zuyderzee, Walfish Bay, phosphate H₂S, O₂, salinity, N.W. Polder, Zurich See (see Minder, 1943).¹⁷²

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Definition of biocoenosis.¹⁷³

Plants.

Proverbs 31:25 The ants are a people not strong, yet they prepare their meat in summer.

Ant-plant!

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