Text box 7.1 – Baas Becking notes made prior to writing the manuscript

“Closed universes”, J. Beauverie and S. Monchal. Comptes Rendus, 195 (1932).1

Coenobiosis Derx, symbiose, metabiose, pseudometabiosis.

Antagonism, L. v. Luyck.

M. Kiese, Klinische Wochenschrift 22, 505, 1943. Penicillin!

Abraham, Cole and Porter, Raistrick.2

Unicellular, all potencies (possibilities) together. Possibilities spread! Is it all redox? Is it all exchange of substances? (See Fig. 7.1).

Maximal to n potencies.

“Autotrophy”, antagonism, serology!

“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.

Text box 7.2 – Baas Becking notes made prior to writing the manuscript

Ergones. (v. Euler, Arch. Chem. Mineral. Geol. Bd. 11a, 12, 1934).19

G. Harmsen in Aerob. Cell decomposition finds many “bios”-like substances, “pure cultures are often not even viable on the ordinary synthetic media”.20Prepared media.

Die Wuchstoffe in der Mikrobiologie. A. Janke.21

7.2.1 Introduction

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.

7.2.2 Carotenoid

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).

7.2.3 Vitamin B complex, aneurine, nicotinic acid

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

[Baas Becking inserted Fig. 7.5.]

Aneurin, Co carboxylase, thiasole.

7.2.4 Bios complex biotin, pantothenic acid, inositol30

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

7.2.5 Ascorbic acid and dienole

Ascorbic acid C6H6O8,


Reductive acid (von Euler),


dioxymalic acid,




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

7.2.6 Steroles

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).

7.2.7 Auxins

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

7.2.8 Phyllochitins

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.

7.2.9 Other vitamins and possible ergones

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

7.2.10 Summary and conclusions

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!

7.3.1 Introduction

[Baas Becking left this section blank.]

7.3.2 Temperature series according to Miehe47

[Baas Becking left this section blank.]

7.3.3 Epiphytes

[Baas Becking left this section blank.]

7.3.4 Epizoa

[Baas Becking left this section blank.]

7.3.5 Summary and conclusions

[Baas Becking left this section blank.]

7.4.1 Introduction

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

7.4.2 Histosymbiosis and ontogeny

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

7.4.2. a Tissues

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.

7.4.2.b Hormones as maintenance

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.

7.4.3 Gamosymbiosis

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.

7.4.4 Phylosymbiosis57

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

7.4.5 Embryosymbiosis

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.

7.4.5. a Diploid dependent upon haploid

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.

7.4.5.b Haploid dependent upon diploid

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.

7.4.5.c Diploid upon diploid and upon triploid

(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.

7.4.6 Politeia

(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.

7.4.7 Summary and conclusions

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.

Text box 7.3 – Baas Becking notes made prior to writing the manuscript

“Zooxanthellae” (chrysudella) [?],64 in Foraminafera and Radiolaria,

“Chlorella”,65 in Paramoecium aurelia,66Stentor amethystine.67

Hypermastigina, termites and woodroach, Cryptocercus,

Cleveland demonstrated digestion of cellulose, see Fig. 129-131.68

Trichonympha, Leptospironympha from stomach ruminants 1937.

Kudo, Fig 167, 19.69

Charles Nicolle, Naissance, Vie et Mort des Maladies Infectieuses.70

Noel Bernard, 1909.

Parasitium, commersalism.

Cyclic Ardisia, Lolium, Calluna. MIMICRY.

Reinfectuous legumes, Hippophae, Mycetomes, Gunnera, Lichens, Cycas, Podocarpus, Zooxanthellae, Termites, Azolla, Lemna, Livenruts, Orchids, Lycopods, Eusporangiates, artificial symbiosis, catotrophic mycorrhi, strawberry Antlers.

7.5.1 Introduction

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).

7.5.2 Cyclic symbiosis

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

[Baas Becking inserted Fig. 7.9.]

7.5.3 Lolium

Mej J. van Roon.76

7.5.4 Calluna


7.5.5 Non cyclic symbiosis Legume

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

7.5.6 Hippophae, Alnus, Elaeagnus79

[Baas Becking inserted Fig. 7.10.]

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

7.5.7 Gunnera

[Baas Becking inserted Fig. 7.11.]

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

7.5.8 Cycads

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.

7.5.9 Lichens

[Baas Becking inserted Fig. 7.13.]

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

7.5.10 Endothophic mycorrhiza


Bernard (1909), Rumpff (1741-1750).85

7.5.11 Lycopods, Eusporangiates

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

7.5.12 Lemna, Liverwort


Straszburger. 87

7.5.13 Zoosymbiosis, Protozoa

[Baas Becking left this section blank.]

7.5.14 Zooxanthellae

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

7.5.15 Mycetomes93

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

7.5.16 Termites

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.

7.5.17 Herbivorous mammals

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

7.5.18 Higher forms of symbiosis


7.5.19 Artificial symbiosis

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

7.5.20 Mimicry

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.


Birds eggs.

Winter pelts in mammals.

Geometer caterpillars.

7.5.21 Summary and conclusions

[Baas Becking inserted Fig. 7.14.]

Text box 7.4 – Baas Becking notes made prior to writing the manuscript

H. [= G.W.] Harmsen diss. many instances in aerobic cell describe metabiosis.102 See Derx (1943). Silicaplate + bluegreens, later Azotobacter, see Dooren de Jong, Funke.103

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

7.6.1 Introduction

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.

7.6.2 Saprophytism

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.

7.6.3 Pütters hypothesis109

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.

7.6.4 Succedaneous cultures

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.

7.6.5 “Bodemmüdigkeit”

(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.

7.6.6 Spoilage of water and biological purification

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.

7.6.7 The succession on sterile soil: the lava problem

(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.

7.6.8 Plagues, parasites, epiparasites (see Lotka, 1924)122

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

7.7.1 Introduction

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.

7.7.2 Phage and serology

[Baas Becking left this section blank.]

7.7.3 Differentiation

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.

7.7.4 Animal protozoa

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

7.7.5 Round worms

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.

7.7.6 Insects

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.


Mosquitos, Stegomyia, Anopheles.

7.7.7 Bacteria

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

7.7.8 Other cryptogames

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.

7.7.9 Higher plants




Text box 7.5 – Baas Becking notes made prior to writing the manuscript

Feldmann (1930) collected sulphur in Turkmania brought back to Leningrad it burned holes in cloths and paper labels were destroyed; sulphuric acid occurred up to 5.16 % (desert Karakorum).140

7.8.1 Component’s life cycle and milieu cycle

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.

7.8.2 Cyclic phenomena in general

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.

7.8.3 Vegetation cycles and cyclic phenomena in agriculture

(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.

7.8.4 Annual and diurnal cycle, tides

[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!

7.8.5 The cycles of elements (compounds)

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)


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.]

7.8.6 Carbon cycle145

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 C4- 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:

7.8.7 Nitrogen cycle147

[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 N3- 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.

7.8.8 Sulphur cycle149

[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 SSO42, 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 S2- 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.

7.8.9 Summary and conclusions

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.

7.9.1 Introduction

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.

7.9.2 “Closed universes”

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.

7.9.3 The aquarium at Amsterdam

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.

7.9.4 The black mud

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+ + S2- → 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.]

7.9.5 Thermal springs

  1. Amoebae.

  2. Diatoms.

  3. Flies.

  4. Beetles.

  5. Bluegreens.

  6. Sulphur bacteria.

7.9.6 Ocean brine

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

  • b) Artemia.169

  • c) DunaliellaEphydra.

Artificial brine (Boekelo).170

7.9.7 Desert brine

a) LochmiopsisBrachionus.

b) DunaliellaEphydra.


7.9.8 Heatherpool


7.9.9 Mono lake

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


7.9.10 Caves

[Baas Becking left this section blank.]

7.9.11 Disturbed equilibria

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

[Crossed out: 12 Deep Sea.]

7.9.12 Meadow land

[Baas Becking left this section blank.]

7.9.13 Sea clay

[Baas Becking left this section blank.]

7.9.14 Higher organisms

Definition of biocoenosis.173


7.9.15 Higher organised animals

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


[Baas Becking left this section blank.]


   Jean Beauverie (1874-1938), French botanist and mycologist. The reference is to Beauverie and Monchal (1932). Beauverie and Monchal demonstrated that in a closed system plants could live in light four years. This seemed possible due to the equilibrium of gasses exchanged by respiration and assimilation by chlorophyl. No fruitation was observed. The growth was dependent on the amount of water available in the system.
   The references to antibiotic research of A.P. Abraham, M. Cole, C. Porter and R Raistrick, were taken by Baas Becking from Manfred Kiese (1943). Chemotherapie mit Antibakteriellen Stoffen aus Niederen Pilzen und Bakterien. Kiese abstracted research about the ‘wonder drug’ penicillin in the UK and USA from various english language papers on penicillin published in the period 1940-1943. These were unknown to Dutch scientists. Probably Henri Derx informed Baas Becking, when he visited him in the Utrecht prison June 25, 1944, about Kiese’s paper from his contacts with the Delft NG and SF [Netherlands Yeast and Spirits Factory] laboratory and the microbiologists Adrianus Petrus Struyk, a graduate from the Kluyver’s Delft Technical University Laboratory. [Possibly the name ‘L.v. Luyck’ in Baas Beckings note is an error for A.P. Struyk.] Struyk produced in 1944 internal reports for the production of the antibacterial substance produced by Penicillium baculatum, the ‘Dutch penicillin.’
See Burns and Van Dijck (2002), Burns (2005, 2009).
   Ketoacids are organic compounds that contain a carboxylic acid group and a ketone group.
   Georg Franz Knoop (1875-1946), German biochemist. Alongside Hans Adolf Krebs and Carl Martius, he clarified the reaction sequence of the citric acid cycle in 1937. Baas Becking referred to Martius and Knoop (1937), who according to Hans A. Krebs in his 1953 Nobel Prize Lecture:
elucidated the fate of citrate when undergoing oxidation in biological material. Whilst it has long been known that citrate can be oxidated in plants, animals and microorganisms, the intermediary steps remained obscure until Martius and Knoop discovered 𝛼-ketoglutarate as a product of citrate oxidation.
   Reference to Bergmann (1934).
   Probably a reference to Barend Coenraad Petrus Jansen (1884-1962), Professor of Physiological Chemistry University Amsterdam.
   In the 1953 version of Geobiology Baas Becking remarked (p. 647):
For Northwestern Europe forest is climax. Heather protected from sheep will revert to pine, birch forest in Holland. Barbed wire, as a military obstacle protecting certain areas from grazing, will result in the development of the alder. Dunes, protected from man and from rabbits, will show a spontaneous reforestation, the nature of which is dependent upon the calcium carbonate content of the sand. The climax itself is recognised by ecologists to be only a “pause in the eternal change of the vegetation” (Braun Blanquet) and the greatest factor which disturbs the climax is man. The landscape of the largest part of this earth is now definitely “anthropogenic landscape”, from grainfields to tea plantations and teak forests. They have their own climax, or rather climaxes. One of them is the weed patch and the other is the desert. Changes in the milieu are of particular interest in the N.W. European series; sea dune, inland dune, heather, where a progressive “decalcification” of the soil, due to leaching by rain, will give rise to various successions often in close proximity to one another. In this ‘labour shy’ era unwanted successions have given rise to vast ‘green deserts.’
Alder thicket is a minerotrophic wetland community dominated by tall shrubs, especially speckled alder (Alnus incana).
   ‘Succedaneous’ means ‘coming after of replacing something else’. In the 1953 manuscript Baas Becking used ‘successive’ instead of ‘succedaneous.’
   Derx (1947) published about metabiosis. Baas Becking, Kaplan and Moore (1960, p. 266) referred to Derx (1947) metabiosis as “an important concept […] As marchin a play, there is a sequence and a constant regrouping of the actors.”They described the processes, which “seem, at first glance, only biological. But, apart from the actors, there are also abiological elements in the play. The metabiosis is of a mixed nature.”
See for metabiosis Sections 7.6 and 7.8.1.
   The primary photosynthetic tissue, the palisade mesophyll, is located on the upper side of the blade or lamina of the leaf.
   ‘Phytonic’ means in botany ‘relating to a phyton.’ A ‘phyton’ is the smallest unit of a plant that can grow into an entire plant.
   See for ‘synevolution’ Section 5, Adaptation and Section 7, Phylosynthesis.
In the 1953 version of Geobiology Baas Becking further elaborated the concepts under the heading Synevolution (p. 677-687). After a quote for A.J. Kluyvers address at the International Congress for Microbiology, Copenhagen July 20-26, 1947, he described his concept of synevolution:
In his Copenhagen address, 1947, A.J. Kluyver states:
“One gets the impression that at all levels of evolution one has to do with living substance that is essentially the same. But during the trend of evolution this living substance has started to lose its independence, the losses in synthetic ability increasing with increasing morphological complexity and increasing adaption to a heterotrophic life. Yet the main lines of metabolism have been maintained, the ultimate requirements have remained unchanged. The difference between the various physiological groups of organisms are not to be found in differences in the fundamental constitution of their living substance, but just in the different ways in which the latter originates. Whether an organism itself synthesises the active groups of its enzyme systems, or has to find these groups ready made in its medium is only of secondary importance; the final result being the same in both cases.”
While serology has taught us that the above statement cannot be taken too literally (as there is ample evidence of specific chemical differentiation, particularly on the protein level) Kluyver has given a lucid summary not only of the present status of our knowledge but also, indirectly, of the future task or biology. Suppose that Neurospora and other, similar, versatile microbes will yield, in the near future, a ‘grand tableau’ of biochemistry, genetics has to shift to morphogenetics and (with the help of physiology) to experimental morphology, to the discipline or morphogenesis. For there lies the mystery of the Darwinian phase or evolution. Lotka [Elements of Physical Biology], already in 1924, has had a clear conception of this attribute of evolution, when he named one of his chapters Evolution Conceived as a Redistribution. He states moreover, (p. 277), that the concept of evolution, to serve us in its full utility, must be applied, not to an individual species, but to groups of species which evolve in mutual interdependence; and, further, to the system as a whole, of which such groups form inseparable part. For to all of the potentialities packed together in a single bacterial cell we have to add (biochemically speaking) almost nothing to arrive at the potentialities of the higher organism, but the great difference lies in the distribution of these potentialities over a range of specialised, but correlated, cells. It may be that many chemical possibilities have been lost on the wayside.
By the term “synevolution” we do not mean that the “completely outfitted” plant cell is, in itself, in origin, a heterosymbiotic creation. Famintzin, in 1907, has claimed that the complete plant cell contained zooxanthellae (plastids), nucleus and protoplasm being an amoeba, while the “microsomes” (mitochondria) represented bacteria. Wallin (1927) is unable to see the difference between intracellular mitochondria and bacteria. No doubt other similar speculations could be traced in the literature. However, as there is not the slightest experimental evidence for these assumptions, it seems that, at the base of the morphological phase of evolution, we should consider a rather large number of different cell types, at different stages of biochemical perfection, some of which gave rise to more differentiated offspring. In this relation the rather unique position of the bacteria, actinomycetes and bluegreens is apparent. They are in some cases, biochemically near “perfect” but the lack of nucleus and (consequently?) the lack of sexual reproduction seems to preclude further morphological development. In more than one. respect they have to be considered as stagnant groups. It would be tempting to compare the phylogenetical, as well as the ontogenetical differentiation with a series of loss mutations as met with in Neurospora or in Chlamydomonas.
Certain organisms, apparently, lack a whole packet of genes. It seems trivial to call these relations synevoluistic, however. The ruminants before their appearance, presuppose the existence or grass. If […] right, the grass presupposes the existence of fungi, and fungi cannot live on their own, their existence presupposes an autotroph. Moreover, the cow could not digest the grass without bacteria and, maybe, ciliates. With this example we are right in the middle of synevoluistic thought, and we may proceed to propose the thesis that flowering plants presuppose the presence of insects, and vice versa. This synevolution must have happened in Mesozoic times.
See also Sapp, Carrapiço and Zolotonosoc (2002).
Ivan E. Wallin (1883-1969), American biologist, claimed that mitochondria were symbiotic bacteria. See Sapp (1994, p.46-51 and 113-116); Sapp (2003, p. 240-241).
   Reference to Bjørn Føyn (1898-1985), Norwegian zoologist, worked as research fellow from 1932 to 1937 with Rolf Nordhagen (1894-1979) at the Bergen Museum. He criticised racial biology as practised in Nazi Germany. During WWII in German occupied Norway he was incarcerated some time in Nazi concentration camps.
Baas Becking repeatedly referred to Føyn. Because his remarks are very personal observations, the references are quoted in full. Baas Becking (1946b) remarked:
Ophrys apifera is an orchid whose flowers, both in shape and scent, imitate the female of a digging [= digger] wasp (Føyn). The imitation is perfect in almost every detail, as photographs show. Even the scent emitted by the female seems to be identical with the odour of the flower. In phylogeny as well as in ontogeny, therefore, we meet the statistically unpredictable, the ‘case unique’, the element of what we are prone to call ‘choice’ or else a directing power superimposed upon the otherwise recurrent events which, together we call ontogeny.
In a lecture given in Sydney, Baas Becking (1951b), Forgotten Biology, he again used this example.
There is a small terrestrial orchid, occurring in Marocco and in Western Europe, called Ophrys apifera, the bee orchid. A Norwegian botanist, Føyn, a pupil of Nordhagen, has made the observation, now twenty years ago, that only the lower flowers at the stalk set fruit, the upper flowers remaining sterile. It appeared that the oldest flowers, were pollinated by the males of a digger wasp, the wasps being attracted both by visual and olfactory stimuli. After the females of the wasps emerged from the cocoons later in the season, the males lost interest in the flowers and the later flowers remained, therefore unfertilised. The attraction of the males proved to be a sexual attraction, the flower of the Ophrys imitating the female digger wasp to such an extent, that, as Føyn convincingly showed by photographs, even the slightest spot on the wings and on the body of the female wasp is reproduced by the orchid. Furthermore, it could be shown that the scent of the orchid, if not identical, is closely related to the scent given off by the female wasp. It seems futile to try to “explain” this mechanism. There are about 30,000 species of orchids, and 5,000 species of digger wasps. The chance that the observed mimesis could be developed by means known to us, even such powerful means as natural selection, is nil. The consequences of this realisation are far reaching. In the first place it gave me the impression that, as a scientist, I am on a big game hunt, equipped with a pea rifle, with toy tools, asked to explain the intricacies of a complicated machine. I might hit a small bird with the pea rifle, I might succeed to unscrew one nut from the complicated machine with my toy wrench but, if I remain honest, I will realise the futility of my efforts.
In the 1953 version of Geobiology Baas Becking described the symbiosis between orchids and digger wasps (p. 597):
The orchid Ophrys is pollinated by a digger wasp. The orchid Cryptostylis is pollinated by an Ichneumonid. Both orchids are able to secrete a substance identical with, or closely related to, the substance secreted by the respective female wasps, causing a strong taxi in the male. Here the definite organic compound enters into this ontomimetic symbiosis [= symbiosis via mimesis of the ontology], but the story is by no means ended, and the phenomenon accounted for. By further specification of this substance, we would be as much in the dark as to the origin of this mechanism, which mechanism, for a long time to come, will defy causal explanation. For, with the specificity of this “attractor substance” we have only touched upon one phase of the problem. Cryptostylis flowers crudely resemble, and certain Ophrid flowers closely resemble the females, the odour of which they imitate. Here the morphological factors as well as the chemical factors blend to create an entity, and this entity is life itself. And I am not ashamed to use the word ‘beauty’ in this connection.
   Max Hartmann (1876-1962), German biologist. Author of studies about sexuality such as: Hartman (1940). Hartmann was the PhD advisor of Franz Moewus (1908-1959), the controversial German scientist.
The investigations on sex hormones in the green algae Chlamydomonas carried out by Moewus, Kuhn and collaborators at Heidelberg have been extensively reviewed and all attempts to repeat critical features of the work failed. See Sapp (1990); Deichman (2001, p. 329-336).
In the 1953 manuscript of Geobiology Baas Becking referred on several pages to the work of Moewus. In the 1953 manuscript of Geobiology he referred to Moewus (p. 106-107). Baas Becking met Franz Moewus when Moewus and his wife Liselotte were working as Timbrol Fellows in the Botany Department of Sydney University (November 1951-1953):
If one has had the privilege, like the author, to see Frans Moewus performing Chlamydomonas, one is amazed at the complexity of the behaviour of these unicellular algae and of the physical and chemical factors controlling their various coordination’s with the environment. For these, and similar, so called “simple” organisms the rules governing the behaviour, growth and development are already beyond us. Even if we could account for all of the processes of differentiation and of coordination in the development of the higher organism, we would be still be unable to integrate them into a rational concept of the organism as a whole, in tune with its environment. And after this riddle there are other mysteries, those of relationship and of descent. The last century has given us the wholehearted cooperation of chemists, physicists, and mathematicians. the mere fact that these workers, so highly successful in their more accessible fields have failed to point out to us even where the riddle lies, has convinced the author that the central problem of life, the understanding of the vital state, is still outside the present realm of the more exact sciences. This does not mean that advances in biology, especially in biochemistry have not been commensurate with those in the more exact sciences, but only that the problem of life is so complex.
The ‘organism as a whole’ concept is a reference to Smuts (1926), Holism and Evolution. See also Section 1.4.
   Hans Spemann (1869-1941), German embryologist. The Spemann-Mangold organiser, also known as the Spemann organiser, is a cluster of cells in the developing embryo of an amphibian that induces development of the central nervous system. Hilde Mangold (1898-1924) was a PhD candidate who conducted the organiser experiment in 1921 under the direction of her graduate advisor, Hans Spemann, at the University of Freiburg in Freiburg, German. Her thesis was published in 1924. Without her consent Spemann had added his name as first author. The discovery of the Spemann-Mangold organiser introduced the concept of induction in embryonic development. On September 4, 1924, Mangold died of severe burns caused from a kitchen explosion in her apartment. Therefore, she did not share the Nobel Prize in Physiology or Medicine awarded to Spemann in 1935.
In the 1953 version of Geobiology Baas Becking remarked (p. 19):
We know next to nothing about morphogenesis itself, but the work of Needham (1931) establishes the main outlines of its chemical basis. Already in 1921 Spemann and, in 1933 Brachet [Belgian biochemist Jean Louis Auguste Brachet, 1909-1988] adumbrated the possible chemical nature of the “organising principle” in ontogeny. Since then, many facts have come to light which have clearly demonstrated the role of specific hormones, enzymes and inhibitors in the development of the organism. It should be stated that already Haberlandt, in his Physiologische Pflanzenanatomie had foreseen this possibility.
See Spemann and Mangold (1924), De Robertis (2009), Sapp (1997), Sapp (2003, p. 176-177).
   Reference to Joseph Needham (1900-1995), British biochemist. Baas Becking referred to his three volume Chemical Embryology (Needham, 1931).
   In the 1953 manuscript of Geobiology Baas Becking discussed ‘Heterosymbiosis’ as a complex of binary combinations between organisms (p. 610-611).
The binary relations between sixteen groups of organisms, yielding 16 x 15/2 or 120 combinations. As the species inside each group will show mutual symbiotic relations at least 136 groups of binary combinations are possible. […] It would require a comprehensive study of the widely scattered literature on subjects as field biology, ethology and ecology to increase the number of relations materially.
In a note he remarked:
Relations between molecules, as governed by mass laws, chemical kinetics (with the inclusion of catalysis) are, of course, much better understood than relations between organisms. The same pertains to celestial bodies. There is a curious analogue, however, with the latter; in the attempts at a mathematical analysis of the relations between two organisms, already considerable difficulty has been encountered (Lotka, 1924), while a three component system gives rise to very complicated relations indeed. Here we meet with an analogue, of the ‘three body problem’ of the astronomer. Pluricomponent systems have proved, as yet, little amenable to mathematical analysis.
   A reference to an antibiotic oxalicum product of Penicillium oxalicum. Penicillium oxalicum produces secalonic acid D, chitinase, oxalic acid, oxaline and β N-acetylglucosaminidase and occurs widespread in food and tropical commodities. This fungus could be used against soil borne diseases like downy mildew of tomatoes. Penicillin and oxaline were at the moment Baas Becking wrote this text not available as medicines. Production of penicillin started when the US entered WW II. During the War, penicillin made a major difference in the number of deaths and amputations caused by infected wounds among Allied forces, saving an estimated 12 %–15 % of lives.
See also Baas Becking’s annotations at the beginning of Section 7 about penicillin research during WWII in Germany and The Netherlands.
   Baas Becking referred to Hans Karl August Simon von Euler-Chelpin (1873-1964), German born Swedish biochemist, won the 1929 Nobel Prize in Chemistry for research on alcoholic fermentation of carbohydrates and the role of enzymes. However, the work on “ergones” was done by his son Ulf Svante von Euler (1905-1983), Swedish physiologist and pharmacologist, who received 1970 Nobel Prize in Physiology and Medicine for his work on neurotransmitters.
   Reference to Harmsen (1946). The research was finished in 1939, but published after WWII. See also Section 3.9.
   Possibly a reference to Janke and Sorgo (1939).
   Baas Becking referred to the act of Molière’s Le Malade Imaginaire (1673) where there is an exchange between the dancing doctors speaking in a fake Latin language (“B.B.’s ‘Board’”) and the chorus:
Le Premier Docteur : Domandabo causam et rationem quare Opium facit dormire.
Le Bachelier [the Bachelor] : […] quia est in eo virtus dormitive, cujus est natura sensus assoupire.
Chorus: Bene, bene, bene, bene respondere: Dignus, dignus est entrare in notro docto corpore.
   ‘In litteris’ means ‘In letters”. Apparently, Henry Derx wrote a letter to Baas Becking when he was in the Utrecht prison. July 25, 1944, he visited him in prison together with T. Niekerk-Blom. They were allowed to speak with him for 25 minutes and gave him two books at that time. Source: NIOD 214 nr 33. Apparently, Baas Becking and Derx discussed the subject of ‘symbiosis’ during this visit, which Derx in his letter [not retraced] further explained.
   Roland Thaxter (1858-1932), American mycologist, renowned for his contribution to the insect parasitic fungi. Thaxter wrote in the 1890s several studies about Myxobacteriaceae in the Botanical Gazette, e.g., On the Myxobacteriaceae, a New Order of Schizomycetes.
   Reference to Pier Andrea Saccardo (1845-1920), Italian botanist and mycologist. Baas Becking quotes from Saccardo’s Sylloge Fungorum omnium Hucusque Cognitorum.
   See for Bassalik also Section 7.1.
   Reference to Carr and Price (1926). The Carr-Price reaction was employed in testing biological materials for vitamin A. The amount of blue colour produced by the reaction of vitamin A with a chloroform solution of antimony trichloride (SbCl3) is proportional to the amount of vitamin A present.
   Endophthalmitis means bacterial or fungal infection inside the eye involving the vitreous and/or aqueous humors.
   Reference to Dr. Joseph Goldberger (1874-1929), Hungarian American epidemiologist who studied the pellagra and demonstrated that pellagra was associated with diets deficient in animal proteins. Pellagra is a disease caused by a lack of vitamin B3. Symptoms include inflamed skin, diarrhoea, dementia and sores in the mouth. The disease is manifested in dogs as ‘black tongue.’ Baas Becking’s son Jan Mattias Baas Becking died of pellagra in 1945 in the Japanese Mariso prisoner camp at south of Macassar (Sulawesi). The Mariso camp housed prisoners of war from June 1944 to August 16, 1945. In that period, 330 people were killed in the camp (out of 1,500 camp residents).
   Vitamin B complex, includes eight vitamins soluble in water.
   Wildiers (1901) discovered that yeast required a special growth factor, which he named “bios”. It proved to be a mixture of essential factors, one of which was biotin. Baas Becking referred to the Liebig-Pasteur dispute on the processes and causes of fermentation. Pasteur supported the idea that fermentation was a biological process, Von Liebig supported the idea that fermentation was a chemical process, discrediting the idea that fermentation could occur due to microscopic organisms. Eduard Büchner uncovered in 1897 the nature of alcoholic fermentation by his discovery that yeasts produce an enzyme that catalyses the chemical fermentation process both inside and outside cells.
Reference to Eduard Büchner (1860-1917), German chemist, winner of the Nobel Prize in Chemistry in 1907. The classical treatises are Büchner (1897a and 1897b).
   Baas Becking referred to the Reichstein-Grüssner process designed for vitamin C production on industrial scale in 1933. D sorbitol is converted to L ascorbic acid using a fermentation step (bioconversion of D sorbitol to L sorbose by Gluconobacter oxydans) and several chemical steps (from L sorbose to L ascorbic acid).
   In this section on Ascorbic acid and Dienol, Baas Becking referred to H. von Euler-Chelpin’s work on oxidation and reduction of ascorbic acid.
   Reference to Giroud (1938).
   Reference to PhD thesis A. Quispel (1943, p. 478-479).
   Stoll (1937. The cardiac glycosides are a group of plant materials arbitrarily so named because of their specific “digitalis-like” effect on the heart muscle. Chemically, the cardiac glycosides are steroid derivates.
   Reference to Adolf Friedrich Johann Butenandt (1903-1995), German biochemist who was awarded Nobel Prize in Chemistry in 1939 for “his work on sex hormones.” He initially rejected the award in accordance with government policy, but accepted it in 1949. Butenandt’s involvement with the Nazi regime and various themes of research led to criticism after the war, and even after his death the exact nature of his political orientation during the Nazi era has never been fully resolved.
See Trunk (2006), Deichmann (1996, 2001).
Also reference to Leopold Ružička (1887-1976), a Croatian-Swiss scientist and joint winner of the 1939 Nobel Prize in Chemistry “for his work on polymethylenes and higher terpenes […] including the first chemical synthesis of male sex hormones.” From 1927 till 1930 he occupied the Chair of Organic Chemistry at Utrecht University.
   The heteroauxine is indole-3-acetic acid (IAA), was isolated in 1934 by Fritz Kögl (1897-1959) and co-workers from human urine; they named the auxin ‘hetero-auxin.’ In the early 1930 Kögl and co-workers had isolated non-indole auxins: auxin a (auxentriolic acid) and auxin b (auxenolic acid) also from human urine, which they considered as the growth hormone auxin in plants. In the above text Baas Becking therefore characterised these substances as the ‘auxins proper.’ Nowadays however, IAA is accepted as the plant growth auxin, because auxin a and auxin b were found to be not natural plant products and IAA has since been isolated from numerous plant species and has shown to be ubiquitous in the plant kingdom. The auxin a and auxin b issue is nowadays considered to be a fake of Kögl’s assistant Hanni Erxleben (1903-2001), although Kögl was possibly also implied.
See Haissig and Davis (1994), Arteca (1995, p. 1-12, Discovery of Plant Growth Substances); Deichmann (2001, p. 339-342), Wildman (1997).
Rhizopon A tablets containing hetero-auxine were produced in the 1930s by the N.V. Amsterdamsche Chininefebriek and used in oculation in plantculture.
   Reference to PhD thesis Went (1928).
   Reference to Koningsberger and Verkaaik (1938).
   Reference to Herbert McLean Evans (1882-1971), who in 1922 at Berkeley along with Katharine Scott Bishop (1889-1975) co-discovered vitamin E. (Evans and Bishop, 1922). They found that the dietary factor, essential for reproduction in rats, is fat soluble and present in green leaves (lettuce), and occurs in special high concentration in wheat germ.
   Reference to the work of George Oswald Burr and Mildred Burr, who discovered in meticulous analysis of rats fed special diets that fatty acids were critical to health. If fatty acids were missing in the diet, a deficiency syndrome ensued that often led to death. The Burrs identified linoleic acid as an essential fatty acid.
   Reference to William Robert Fearon (1892-1959), Irish politician and Professor of Biochemistry at the University of Dublin. He published about biochemical colour tests. In 1925 about Colour Reactions Associated with Vitamin A.
   In The 1953 manuscript of Geobiology (Baas Becking, 1953a, p. 592 and 593), Baas Becking used the following classification of symbiosis, which is a further enlarged version compared with that in the 1944 manuscript of Geobiology:
KTENOSIS. The killing of the other, for food, in defence, or for other reasons.
ANTAGONISM. The exertion of an adverse influence, which need not result in another’s death.
PARASITISM. To subsist upon another without the primary object of killing.
HELOTISM. To exploit another.
MUTUALISM. To benefit by another’s activities.
COMMENSALISM. “To feed at a common trough”.
SAPROBISM. To subsist on the refuse of others.
MIMISIS. To imitate another, or an inanimate object.
METABIOSIS. The succession of organisms.
The scale of complexities of the substrate could be stated as follows:
CYTOSYMBIOSIS. example; phagocytosis.
HISTOSYMBIOSIS. “organiser”, hormones in correlative metabolism.
GAMOSYMBIOSIS. sex substances in flagellates, ferns, flowers, animals
ONTOSYMBIOSIS. this is the relation between individuals. Example; domestic plants, bacteria in the rumen of the cow.
OIKOSYMBIOSIS. relations of individuals in a community.
   Reference to Fröschel and Funke (1939), Funke (1942).
   Chorism or chorisis is multiplication or dispersal of botanical elements, in the text meant by birds and by ants.
   Reference to Miehe (1907) and Miehe (1930), see also Section 6.5.
   In the 1953 version of Geobiology Baas Becking described autosymbiosis as follows (p. 156-158):
Modern cytology has painted the fascinating and complex picture of the continuity of the innermost, particulate expression of the specific individuality, localised in the nucleo-proteins in the confines of the nucleus. This is, in the space-time continuum, the “germ track”, the “Keimbahn” of the older authors. Sexual differentiation, a specific form of symbiosis (which may be called gamosymbiosis) we know to be a powerful agent in evolution.
With this symbiosis, resulting from differentiation within the organism (an autosymbiosis) the possibility for an endless variation in pattern became possible. The offspring of a single cell gives rise to an organism, the cells of which are mutually dependent. [Baas Becking inserted the remark: like the halt and the blind.] Cells became specialised and took over single functions, where the autarchic unicellular ancestor could perform all of these functions. The developmental pattern was dictated by the specificity of the nuclear matter, by the chromatin.
Unlike in chemistry where a certain molecule under certain environmental conditions will give rise to a certain crystal lattice, even the given unchanged nuclear structure will give rise to beings while similar in shape, in differentiation and in behaviour, show variation on a common theme. Organisms show variability. And it is this variability that was recognised by Darwin as a second potent factor in evolution. While the nuclear structure, seat of heredity characters is, in principle, a fixed entity, the soma, the protoplasm, the vegetative characteristics are variable, and this variability is influenced by the external environment by the “milieu exterieur.” However, the above concept would yield only a static structure, unable to account for hereditary changes necessary for evolution.
While we are agreed that such changes must occur, the main question centres on the relative importance of the outer and of the inner world and the interplay between them. Before we consider this question further; it might be of use to consider the concept of symbiosis more in detail. While highly differentiated morphologically, a so called “higher organism” cannot perform more ‘chemical feats’ than a single algal cell. This morphological differentiation, giving rise to hundreds of different cell types, is an expression of coordinated autosymbiosis. The higher organism is a vital community, a living landscape, a biocoenosis in itself. But only rarely, if ever, it can exist by itself. Apart from its inner coordination, it is equally bound, by many ties, to both the animate and to the inanimate environment. It is only a part of the warp or of the woof of the tissue of life. And, in organic evolution, one organism cannot “move onward” without the necessary readjustment in the whole tissue.
   See notes above.
   Refers to Gaillard (1942). Pieter Johannes Gaillard (1907-1992), in 1947, became Professor in Experimental Histology at Leiden University. He finished his book in 1940 at the beginning of WWII. The manuscript was published without his knowledge in Paris in 1942.
   The parathyroid glands are small endocrine glands in the neck of humans which secrete parathyroid hormones in response to low blood calcium.
   See Section 7.5.15.
   Baas Becking’s did not have a classical education, so his Greek is not reliable. Perhaps he wanted to refer to the Greek word for ‘message’: Mήνυμα.
   Wilhelm Friedrich Philippe Pfeffer (1845-1920), German botanist and plant physiologist. Baas Becking referred to Pfeffer (1884).
Christian Ernst Stahl (1848-1919), German botanist; Baas Becking referred to Stahl (1884).
   The reference is to Kuhn, Moewus and Wendt (1939). The experiments of Moewus and Kuhn have been extensively reviewed and all attempts to repeat critical features of the work failed. Nowadays the results of his studies are considered as fraud. See Sapp (1990), Deichmann (2001, p. 329-336), Sapp (2003, p.154-166).
In the 1953 version of Geobiology (p. 601-605), Baas Becking referred to Moewus and Kuhn in the Section Homoio Symbiosis, Section Carotene and Derivatives. Apparently, he was at that time not informed about the discussion about the validity of Moewus experiments. According to Baas Becking:
Not only the carotene itself, but many of its derivatives are intimately connected with sexual reproduction, particularly derivatives of the glucoside crocin. Particularly through the work of Kuhn, started before the war, interesting and significant facts have come to light. As Moewus (1950a and b) has complemented his early work by cytological and genetical studies (using, as in the Neurospora work, artificially induced mutations), the sexual relations of the green flagellate Chlamydononaseugametos have been shown to be dominated by several specific substances. These substances are formed from precursors by chain reactions, every link in the chain corresponding to one or more genes. Chlamydomonas has 10 chromosomes, and 10 linkage groups have been observed. The picture as given by Moewus in 1950 is less homogeneous as it appeared to be in 1938, as flavonol glucosides and their derivatives happen to play an important role. Moewus distinguishes “gamones” i.e. ergones, determining the attraction between gametes, further “termones” which differentiate the sexes in hermaphroditic strains. He also finds substances causing flagellar motility and flagellar growth, while also an inhibitor of copulation was found.
[….] Every step has been checked by genetical evidence obtained by mutants. Moewus also succeeded in isolating several of the substances involved from the Chlamydomonas cultures. It is interesting to note that, in his work, Moewus makes no reference of the analogous work with Neurospora. Apparently, there has been a parallel development of two, very important, branches of research.
   Reference to Jacques Loeb who took a decided stand against chemotropism of sperm in animals. See U.B. Kaupp (2012) 100 years of sperm chemotaxis.J.Gen. Physiol. 140 (6), p. 583-586.
   Phylosymbiosis can be defined as microbial community relationships that recapitulate the phylogeny of the hosts. Recent findings indicate that mammalian gut microbiome plasticity in response to dietary shifts over both the life span of an individual host and the evolutionary history of a given host species is constrained by host physiological evolution. Therefore, the gut microbiome cannot be considered separately from host physiology when describing host nutritional strategies and the emergence of host dietary niches.
See Amato et al. (2019) and Brooks et al. (2016).
   Noël Pierre Joseph Léon Bernard (1874-1911), French botanist, discoverer of the symbiotic germination of orchid seeds where a soil fungus provides water, mineral nutrients and carbon to the seedling, and compensated for the absence of reserves. Baas Becking referred to Bernard’s 1899 article Sur la Germination du Neottia Nidus-avis. See Selosse, Minasiewics and Boullard (2017).
Joseph Magrou (1883-1951), cousin of Noël Bernard and his co-author of Sur les Mycorrhizes des Pommes de Terre Sauvages (Bernard and Magrou, 1911). After Bernard’s early death in 1911 by tuberculose, his ideas were taken up by his former teacher J. Costantin and J. Magrou.
See also Selosse, Boullard and Richardson (2011), Yam and Arditti (2009).
In the 1953 manuscript of Geobiology Baas Becking quoted Bernard (p. 590) when he discussed the complexity of symbiosis: […] l’état dit de symbiose est en quelque sorte un état de maladie grave et prolongée […]. In Section Synevolution (p. 679-680) he referred to Bernard and Magrou:
Bernard has discovered an important relation between the formation of tubers, or swollen rhizomes, in higher plants on the one hand, and mycorrhizal fungi on the other. Here a fundamental change should occur, according to Bernard, in the organism infected by the fungus. While the formation or tubers in several orchids did not take place without the fungus, in the potato the tuber formation, by centuries of culture, may take place independent of any previous infection. Here something ‘new’ has been created. Apart from the Lamarckistic implications of the theory, there is much to be said for the idea, that part of the morphogeny of the higher plant may be “gall formation” by symbiotes.
The difficulty remains to apply this thought to evolutionary theory. Any symbiosis might imply a “loss” in one or two of the partners. It remains to be seen whether the terrestrial orchids would be able to live without the fungus. If this were the case, the symbiosis would have added something new, the tuberisation. If, however, the plant should appear deficient without its symbiote, if the relation were obligatory, then the combination would be a stable one, acting upon an evolutionary unit. Despite the great mass of fact brought forward to support the theory, however, the direct application to evolutionary theory seems difficult, if not impossible.
   This section is a remarkable plea for Baas Becking’s ideas about ‘synsymbiosis’and ‘synevolution.’ See also Section 5 on, Adaptation and Section 7, Symbiosis and Antagonism.
   ‘Scutellum’ is part of the structure of a barley and rice seed – the modified seed leaf. The scutellum is believed to contain an as yet unidentified protein transporter that facilitates starch movement from the endosperm to the embryo.
   Baas Becking referred to Campbell (1905). Douglas Houghton Campbell (1859-1953) American botanist, specialist in liverworts. In 1921, Baas Becking published his Stanford PhD dissertation (Professor Dr. D.H. Campbell PhD advisor), The Origin of the Vascular Structure in the Genus Botrychium (Baas Becking, 1921b). A descriptive study in plant anatomy in which the embryological development of several species of Botrychium (a fern also known as moonwort) is described.
   Possibly reference to Ezekiel 32:2. Son of man, take up a lamentation for Pharaoh king of Egypt, and say unto him, Thou art like a young lion of the nations, and thou art as a whale in the seas: and thou camest forth with thy rivers, and troubledst the waters with thy feet, and fouledst their rivers.
   Anna Martha Hartsema (1896-1975) defended her doctor’s thesis May 12, 1924 in Utrecht: Over het Ontstaan van Sekundaire Meristemen op de Bladeren vanBegonia Rex (Hartsema, 1926; PhD advisor F.A.F.C. Went). She was well known to Baas Becking as a colleague in the Utrecht Botanical Laboratory. She became a scientific researcher in the Agricultural University Wageningen.
In the manuscript of the 1953 Geobiology Baas Becking referred to Hartsema (p. 685):
The regeneration of a Begonia plant from a single epidermal cell is by no means proved (Hartsema, 1926). In the foliar buds of Bryophyllum, in the gemmae of Marchantia, in the fronds of viviparous ferns. It is a group of cells rather than a single cell which gives rise to a new individual.
   Zooxanthellae is a colloquial term for single celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including corals, jellyfish and nudibranchs.
   Chlorella is a genus of single celled green algae belonging to the division of Chlorophyta.
   Paramecium aurelia, unicellular organism.
   Amethyst deceiver, Laccaria amethystina, a small brightly coloured mushroom that grows in deciduous as well as coniferous forests.
   A number of genera in the group Hypermastigina (flagellates) inhabit the intestines of termites and cockroaches. The Hypermastigina are able to digest cellulose, an ability which generally does not occur among animals.
   Cryptocercus punctulatus Scudden, a cockroach eating wood. Cleveland et al. (1934) found that all Crytocercus had the flagellate Barbanympha in their stomachs and the enzymes cellulase and cellobiase. Further Cryptocercus have two protozoa symbionts Trichonympha and Leptospironympha in their intestines.
See also Kudo (1926).
   Baas Becking inserted this reference to Charles Jules Henry Nicolle (1886-1936), French bacteriologist, winner Nobel Prize in Medicine (1928) for his identification of lice as the transmitter of epidemic typhus. See Section 5.10.13 Epidemiology.
   Reference to Weber-van Bosse (1887). She described Trichophilus welckeri Weber-van-Bosse. This is a subaerial species occurring in the fur of brown-throated, three-toed sloth (Bradypus variegatus; Bradypodidae) and the pale-throated, three -toed sloth (Bradypus tridactylus) which occur from Nicaragua to Brazil. Their long, coarse, thick hair is brownish-grey in colour and takes on a greenish tinge in the rainier season when Trichophilus forms on its back.
   Walter Spies (1895-1942), Russian-born German primitivist painter, composer, musicologist and curator. In 1923, he moved to Java, Indonesia. He lived in Yogyakarta and then in Ubud, Bali. In 1938, he was arrested as part of a crackdown on homosexuals, released in September 1939. As a German national in the Dutch East Indies, he was taken prisoner when WO II broke out. He was one of the victims on the Van Imhoff that transported the German prisoners to Ceylon and that was torpedoed by the Japanese. The German prisoners were not rescued, only the Dutch passengers and crew.
See Ross (2000).
   Adriaan Cornelis Valentin van Bemmel (1908-1990), zoologist, in 1937 assistant curator Zoological Museum Buitenzorg, returned from Indonesia in 1951. Later director of the Rotterdam Blijgaarde Zoo and the Rotterdam Nature History Museum.
   Lieven Ferdinand de Beaufort (1879-1968), Dutch biologist, participated in 1903 in the North New Guinea Expedition. In the 1920s director of the Artis Zoological Museum Amsterdam. Professor in Zoogeography University of Amsterdam.
   The reference is to the studies of Hugo Miehe about Ardisia crispa symbiosis after his sojourn at Buitenzorg and published in 1911, in his Javanische Studien and in Miehe (1917). Ph. De Jongh worked at the Leiden Botanical Laboratory and confirmed the symbiosis Ardisia and bacteria in 1938. Ph.J. de Jongh (1938). The Symbiosis of Ardisia Crispa, PhD Thesis Leiden (Baas Becking PhD advisor).
In the 1953 Geobiology manuscript, Baas Becking referred to their studies on p. 623-624 under the heading Action of Microbiotes on Higher Plants, Foliar Bacterial Symbiosis.:
The bacteroids, present in these nodules were interpreted by him as “proteinaceous glands.” Hugo Miehe, after a sojourn at Buitenzorg published a detailed account of the Ardisia symbiosis in his Javanische Studien (Miehe, 1911). The bacteria are present in the growing point of the plant. Infection of the leaf margins occurs in the hydathodes of the young leaves. Constrictions occur at the margins and in the centre of these contrictions we find the bacterial nodules. In the seed Miehe found bacteria between the embryo and the endosperm. With this study Miehe gave the first example of a cyclic symbiosis between a higher plant and a bacterium, as extraneous infection (as in the Leguminosae) does not take place. In a later study Miehe describes the bacterial component and, in a third study, the bacteria-free Ardisia. The latter is a veritable ‘cripple’, it remains dwarfed, and never flowers. At the Leyden Laboratory, Ph. de Jongh could confirm the results of Miehe (1917). He, moreover, succeeded in following the course of the cyclic symbiosis from seed to seed. The bacterial film, which covers the vegetation points, is enclosed between the carpels. The bacteria finally surround the ovules and are entrapped by the growing integuments at the micropylar region between the two parts of the inner integument. After the (apogamous) development of the embryo, the bacteria adhere to its radical pole, where they remain in a resting, but reversible, state. (this in contrast with the bacteroids in the leaf nodules, which are irreversibly changed). At germination the bacteria. are pushed over the cotyledons, where they infect the axillary bud, while a bacterial mass also infects the terminal bud. Seeds could be made bacteria-free by heating them to 40 °C. The bacteria-free plant, the cripple, has a juvenile appearance. Longitudinal growth stops over the entire plant, the root excepted. The meristematic cells are large, the leaf primordia remain undifferentiated, while the axillary cotyledonary buds proceed to proliferate for years, forming a wart-like gall. The ‘cripple’ contains catalase and peroxidase in the terminal bud while the normal plant seems to lack peroxidase. Application of heteroauxin only caused root formation on stem and petioles, but could not cure the cripple. Cleft grafts of normal on cripple or vice versa showed the root system of the crippled stock to be stimulated by the normal scion. Very young cripples may be artificially infected.
As in the foliar symbiosis of Rubiaceae (Pavetta, Psychotria), fixation of atmospheric nitrogen has been claimed by von Faber (1912-1914), it seems remarkable that the Ardisia system apparently expresses a different relation. de Jongh thinks that a growth promoting substance (not identical with heterauxin) may be produced by the bacteria. As the presence of a high peroxidase seems typical for other ‘dwarfs’, where the dwarfing is caused by mutation of a single gene, it may be that the bacterial film acts as an “oxygen absorber”, reducing the oxygen pressure at the vegetation point. It seems highly significant that R. Bok, at our laboratory, succeeded in inducing growth Ardisia-cripples in oxygen-nitrogen mixtures with a lowered oxygen tension (1941). However, the real nature of the consortium remains unexplained. The relation, while cyclic, remains intercellular.
R. Bok was research assistant ‘without objection from the Governments Treasury’ [‘buiten bezwaar van ‘s Rijks Schatkist’] at the Leiden Botanical Laboratory from March till September 1941. In 1946, she worked on soil optimalisation for Lathyrus culture at the TNO laboratory in Delft. Her 1941 experiments were communicated to the Dutch Academy of Sciences by Baas Becking (Bok, 1941):
These preliminary results seem to give a strong indication that the hypothesis, stated in the work of P.H. de Jongh, according to which the bacterial film on the stem vegetation point of Ardisia crispa lowers the oxygen tension in the stem meristem, contains elements of truth.
   Darnel, ryegrass (Lolium temulentum, L.), has been known since Roman times for the poisonous properties of its grain. It was not, however, until 1898 that the presence of an often considerable layer of hyphæ was discovered just exterior to the aleurone layer of the grain; to the action of this fungus layer the poisonous properties are presumably due. In his PhD thesis Quispel (1943, 1946, p. 419) referred to the research of “fungus symbiosis of Lolium temulentum by J. M. van Roon and Mr. J. v. d. Drift” in the Leiden Botanical Laboratory”.
In 1941, van Roon was an assistant employee at the Leiden Zoological Laboratory. Joop van der Drift worked after WWII at the Instituut voor Toegepast Biologisch Onderzoek in de Natuur (ITBON). He defended his PhD thesis in 1950 in Leiden. He initiated an integrated study on various factors influencing soil life in a small forest of Hackfort in The Netherlands.
   Reference to Mabel Mary Cheveley Rayner (c. 1890-1948), english mycologist, in 1913 Lecturer in Botany, University College, Reading. Rayner thoroughly reviewed academic research into mycorrhizal research.
   Reference to Hellriegel and Wilfarth (1888), Beijerinck (1888), Beijerinck (1918).
‘Fred’ refers to Edwin Broun Fred (1887-1981), American bacteriologist, President University of Wisconsin-Madison. In Geobiologie (1934, p. 85 English edition, 2016), Baas Becking referred to Domogalla and Fred (1926) on solubility of ammonia and nitrate in freshwater.
“Allen” refers to the Ethel K. Allen (1906-2006) and Oscar Nelson Allen (1905-1976), bacteriologists at University of Wisconsin, who published the results of their 45 years of research in 1981: The Leguminosae, a Source Book of Characteristics, Uses, and Nodulation (Allen and Allen, 1981).
“Löhnis” refers to Dr. Marie Petronella Löhnis, a plant pathologist who worked in the 1930s and 1940s on boron deficiency at the Wageningen Landbouwhogeschool (Löhnis, 1941).
Artturi Ilmari Virtanen (1895-1973), Finnish chemist recipient 1945 Nobel Prize in Chemistry.Virtanen worked at the Laboratory of the Foundation for Chemical Research in Helsingfors (Finland) on root nodules of Leguminosa.
   The species mentioned by Baas Becking have nitrogen fixing root nodules. See Hawker and Fraymouth (1950).
   ‘Oven’ probably refers to A. van Oven, a student of Baas Becking in Leiden. Miss Oven participated in the field research of the Leiden Biologist Club in Wijster, autumn 1932, where she demonstrated the presence of bacteria in the air cells of Sphagnum cymbifolium (Baas Becking and Nicolai, 1934). In Geobiology (1953, p. 581) Baas Becking mentioned:
Miss van Oven studying the acid water of a peat bog found active sulphate reduction and anaerobic acid formation by Plectridia [Glucose and cellulose fermenting hammer-shaped or drumstick-shaped bacteria].
   ‘Hansen’ is possibly F.H. Hansen, of the Forest Research Institute, Buitenzorg, Java. In the 1953 manuscript of Geobiology Baas Becking referred on p. 621 to:
The curious algal symbiotes in Cycadeae and in the ‘giant herb’ Gunnera have attracted much attention since Reinke (1873) described the “algal zone” in the root or Cycas. […] In Gunnera (studied by Reinke and later by Miehe) the algal galls (formed by internal infection of the rhizome through the mucilaginous glands) were found to occur always above a root primordium. Just before the war, van Hulssen, at Buitenzorg showed that heterauxin was present in the algal colonies (Gunnera macrophylla) (see Baas Becking, 1947a). The experiments were repeated by Dykshoorn in Leyden with Gunnera manicata, and our earlier results confirmed. It also appeared that the plant contributed minimum substances which influenced favourably with growth of the alga. […] It seems that the Gunnera rhizome is not capable of forming roots without the algal symbiote. This does not exclude the possibility of nitrogen fixation by the alga, but from the relation between the galls and the root formation it appears to be obvious that the chief role of the Nostoc may be its contribution of root-forming substances.
   Baas Becking inspected genera from the order Cycadales. The Cycadaceae are a family of the suborder Cycadineae with one genus Cycas with about 115 species. From the suborder Zamlineae he mentions the genera Stangeria, Dioon, Encephalartos, Macrozamia and Zamia.
   Galestin repeated the experiments of Beijerinck with isolated root nodules of several Legumiosae.
   References to De Bary (1864), De Bary (1879) and Schneider (1898). ‘Jacq’ is Nikolaus Joseph von Jacquin (1727-1817).
   See for Noël Bernard and mycorrhiza of orchids Sapp (1994, p. 13-14) and Selosse, Boullard and Richardson (2011).
The reference to Rumphius is to Georgius Everhardus Rumphius (1627-1702) and his Amboinsche Kruidboek (1741-1750). According to E. Beekman in Rumphius’ Orchids (Beekman, 2003, p. xlv-xlvi):
Rumphius was the first botanist to describe epiphytes and to intuit that these plants were not parasites but that their arboreal hosts were only something of a perch. He was the first to describe orchid seed [….] and seems to have comprehended that orchid seeds are dispersed by the wind. He also clearly understood orchid fruits and was the first to note the presence of pollinia, which have been defined as “more or less coherent masses of pollen.”
See also Beekman (2011), G.E. Rumphius, The Ambonese herbal, which is a wonderful annotated English translation of Rumphius’ botanical work.
   “Treub” referred to Melchior Treub (1851-1910), from 1880-1909 based in the Dutch East Indies as director of the Bogor Botanical Gardens at Buitenzorg. In the period 1884 till 1890 he published about Lycopodium Cernuum in the Annales du Jardin Botanique de Buitenzorg.
H. Bruchman published on the prothallium of Lycopodium. Baas Becking (1921b) referred to Bruchmann (1906).
   Eduard Adolf Strasburger (1844-1912), Polish-German botanist.
   Max Walker de Laubenfels (1894-1960), American spongiologist. The reference is to Laubenfels (1930).
Hilbrand Boschma (1893-1976), Professor of Systematic Zoology, University Leiden and director Museum Natural History Leiden (1933-1958). His chief interest was in systematic zoology particularly that of rhizocephalan crustaceans and stony corals. See Vervoort (1977), Prof. Dr. Hilbrand Boschma 1893-1976.
Jan Verwey (1899-1981), marine biologist and director Zoological Station Den Helder (1931-1965). From 1922 until 1931, he worked as a fishery biologist in the Dutch East Indies.
   Paramecium bursaria, a ciliate in marine and brackish waters, has a mutualistic endosymbiotic relationship with green algae called Zoochlorella. The algae live inside the Paramecium cytoplasm.
   Stentor amethystinus, trumpet animalculus, filter feeding heterotrophic ciliate.
   Baas Becking referred to the phenomenon of coral bleaching which occurs when coral polyps expel algae that live inside their tissues in an endosymbiontic relationship. The cause of bleaching is rising water temperature.
   Possibly a reference to Wolvekamp and Vreede (1941).
   Mycetomes are symbiotic microorganisms, which aid in the digestion of vertebrate blood.
   Reference to Paul Buchner (1886-1978). See also Sapp (1974, pp. 109-11).
   Baas Becking referred to Shelford (1929, p. 322).
   Charles Atwood Kofoid, worked on plankton organisms and protozoa, as Professor of Zoology from 1903 until 1936 at Berkeley and after his retirement in 1936 until his death. His most important contributions to the morphology of the protozoa were made during his association with Olive Swezy (1878-?), zoologist and phycologist at Scripps Institute and University California.
See Goldschmidt (1951).
   Reference to Cleveland (1923), Cleveland et al. (1934).
   The Koala’s digestive system is especially adapted to detoxify the poisonous chemicals in the eucalyptus leaves. Koalas have a special fibre digesting organ called a caecum (200 cm). The caecum contains millions of bacteria which break down the fibre into substances which are easier to absorb. Even so, the Koala is still only able to absorb 25 percent of fibre eaten. Water is also absorbed from the gumleaves, so that Koalas rarely need to drink, although they can do so if necessary, such as in times of drought when the water content of the leaves is reduced.
   In his 1953 version of Geobiology (p. 613), Baas Becking described under the heading MUTUALISM:
Rhodotorula rubra and Mucor ramannianus, Polyporus versicolor and Nematospora gossypii are classical examples of “laboratory symbioses” described, respectively by Schöpfer (1935) and Kögl and Fries (1937). These organisms, each deficient in a certain minimum component, showed good growth when brought together. In the case of Schöpfer’s symbiosis each component could perform a part synthesis of the thiamin necessary for growth, respectively the thiazole and the pyrimidine parts, while Polyporus requires thiamin, provided by Nematospora while, in its turn, it produces the biotin which cannot be synthesised by the Nematospora. So, it seems evident that, in heterosymbiosis as well as in homoiosymbiosis, we deal with “humoral phenomena” and, as in pathology and in physiology, definite solutes are involved.
In the same manuscript (p. 614 and 615) under the heading Excretions of Higher Plants, Mainly Foliar, and their Influence Baas Becking referred to Funke (1943):
G.L. Funke, in 1943, described an interesting effect of the excretions of Artemisia absinthum (which he called “absinthein”) on other plants. Salvia, Nepeta and a number of others were suppressed in their growth when planted within five feet of the absinth. When planted within 40-50 cm from the absinth, they became dwarfed.
   Volucella bombylans occurs in several forms, each of which mimics a species of bumblebee. The two main varieties are Volucella bombylans var. bombylans, showing an orange-red tail, mimicking the red-tailed bumblebee (Bombus lapidaries) and Volucella bombylans var. plumata with a white tail, mimicking the white-tailed bumblebee (Bombus lucorum) and the buff-tailed bumblebee (Bombus terrestris).
   For the colour mimicry of the 65 species of the Dasymutilla velvet ants, see Funke (1943).
   See for G.W. Harmsen Sections 3.9. and 4.8.
   Dr. Louis Edmond den Dooren de Jong (1897-1972), bacteriologist, pupil of Beijerinck. Baas Becking referred to Den Dooren de Jong (1938).
   Henri Derx (1947) published about symbiosis and mentioned Baas Becking’s unpublished views for which he referred to the PhD thesis of Anton Quispel (1943/1946).
   Probably a reference to Giroud (1938).
Humans and other primates have lost the ability to synthesise ascorbic acid (ASC; vitamin C). Unlike primates, mice and rats are able to synthesise ASC. See Gabbay et al. (2010).
   C.A.J.A. Oudemans (1825-1906), Professor of Medicine, Botany and Pharmacology University Amsterdam 1859-1896.
   Reference to Lindner (1888), Die Sarcina-organismen der Gärunggewerbe. ‘Gärunggewerbe’ are fermentation techniques.
   See for Thaxter. Section 7.2.1.
   In the 1934 Geobiologie Baas Becking discussed Pütter hypothesis (p. 85 and 86, English edition). His reference was to Pütter (1911). According to H.W. Harvey (1928, p. 29):
The part played by the traces of filter passing organic matter, occurring in seawater, upon the physiological processes of marine animals in nature, is by no means clear. Pütter (1909) has advanced a theory that they absorb much of their nutriment in the form of dissolved organic matter, but the theory has not been generally accepted on the ground that much subsequent experimental evidence has failed to show any such absorption, and that there is plant life sufficient to support them. The capability of higher marine organisms to obtain nutriment by absorbing dissolved organic matter themselves is not definitely proved or disproved. That marine bacteria and possibly many protozoa utilise these dissolved substances and keep their concentration low in the open sea is probable, if not certain.
Harvey (1928) also referred to Pütter (1909). According to Korringa (1949), the controversy ‘once argued ardently’ by Pütter, whether or not dissolved organic matter can be used as food by marine organisms, is generally thought to be settled with Krogh’s conclusion (1931): ‘There is no convincing evidence that any animal takes up dissolved organic substance from natural water in any significant amount.’
See Krogh (1931), Dissolved Substances as Food of Aquatic Organisms.
   Reference to Birge and Juday (1911). Also quoted in Baas Becking Geobiologie (1934). The reference is to Edward Asahel Birge (1851-1950) and Chancey Juday (1871-1944), pioneers of North American limnology.
   See Kuhn, Moewus and Wendt (1939) and Moewus (1950a and b) and Section 7.4.3. The reference is to the sex determining substances found in Chlamydomonas eugametis. They found that besides ‘Gamonen’ also another sex determining substance, ‘Termone’ was excreted by the gameten. The male determining was oxaldehyde of Safran, the female determining was isorhamnetin. Both substances were still active in extraordinary dilution of about one molecule per cell.
Nowadays the results of Moewus studies are considered as fraud.
   See for G.W. Harmsen above.
   Baas Becking referred to the Wageningen microbiologist Nicolaas Louis Söhngen (1878-1934) and his research on “bodemmoeheid”.
   Refers to publications of Félix Hubert d’Herelle (1873-1949), French Canadian microbiologist and Frederick William Twort (1877-1950), English bacteriologist:
D’Herelle’s short note in the Comptes Rendus in 1917 truly represents what can be recognised as the discovery of bacteriophage (D’Herelle, 1917). While an earlier report in 1915 by F.W. Twort certainly described a phenomenon (Twort, 1915), called “glassy transformation” (of bacterial colonies on agar) and “transmissible lysis”, that was caused by bacteriophage in his cultures, Twort failed to interpret his observations in a way that encompassed the concept of virus, of intracellular parasitism, or of serial reproduction of an infectious agent, all of which d’Herelle proposed with clarity and experimental support in his short first note of 1917.
See Summers (2016) and Duckworth (1976).
   Charles A. Kofoid (1865-1947) American zoologist. The reference is to Kofoid (1903).
   Reference to Dr. Neeltje Louwrina Wibaut-Isebree Moens (1884-1965), who published in 1916 about the quality of surface water in and near Amsterdam (Wibaut-Isebree Moens, 1916).
   Willem Marius Docters van Leeuwen (1880-1960), Dutch botanist and entomologist. Docters van Leeuwen (1936) wrote Krakatau 1883-1933 in answer to Backer (1929), The Problem of Krakatao, as seen by a Botanist. For Cornelis Andries Backer (1874-1963), Dutch botanist. See for the Krakatau Problem Section 4.3.7a.
   According to Quispel (1938) “Geiseltallack was used by geologists to make casts of fossils etc.
In the 1953 version of Geobiology Baas Becking described the method as follows (p. 290):
Systematic measurements of surface tension of natural waters have not been carried out as far as the author is aware. Especially after a seasonal water bloom of bluegreens or a seasonal development of diatoms and Peridinians, such changes might be expected. The study of the water surface in situ has become possible by means of the spreading of solutions of plastics. (B.B., Java 1939, unpublished). When a thin brass ring is dipped in a solution of “Geisselthal-lack” and this ring (diameter ±5 cm), suspended by three threads, is carefully lowered on to the water surface, the area enclosed within the wire will be covered with a thin film in a fraction of a second. Then a glass plate may be placed under the film, which is lifted from the water. It separates quite easily from the brass ring and may be used in direct microscopy. In this way bacterial and algal colonies may be observed in situ.
   Protonema is a thread-like chain of cells that forms the earliest stage (Haploid phase) of the life of mosses.
   See for Funke (1943), Blytt-Sernander and Braun-Blanquet, Section 1.2.B.
   Reference to van Dieren (1934). See also last section, Section 6.
   Baas Becking mentioned A.J. Lotka also as a reference to epidemiology Section 5.9.13.
   Reference to A.F.H. Besemer, botanist, studied in Leiden (until May 7, 1940), in 1941 to Wageningen as a plant pathologist. After WWII Extraordinary Professor in Phytopharmacy Wageningen. Baas Becking possibly referred to Besemer (1942), Die Verbreitung und Regulierung der Diprion Pini-Kalamität in den Niederlanden in den Jahren 1938-1941 (PhD thesis Utrecht).
See also Besemer (1984).
   Reference to Schure (1935a and 1935b). After WWII Petronella Sophia Joanna Schure did research on “Kresek”, a bacterial disease of rice, at the Agricultural Station of the Botanical Garden at Buitenzorg.
   Reference to Woodruff (1914 and 1921). By using natural pond water Woodruff was able to preserve the culture of Paramecium for several years. See also Darby (1930).
Giardia intestinalis, also known as Guiardia lamblia. First likely description in 1681 by Antonie van Leeuwenhoeck.
   Probably Heligmosomoides polygyrus, previously named Nematospiroides dubius, a naturally occurring intestinal roundworm of rodents.
   Present name Enterobius vermicularis.
   Phthirus [or Pthirus] inguinalis parasite of the human uro-genital tract, usually named Pthirus pubis.
   Head louse Pediculus humanus capitis.
   Yersinia pestis is a bacterium that can provoke septicaemic, pneumonic and bubonic plague in humans. The pathogen can be transmitted by infectious droplets or by contact with contaminated fluid or tissue or indirectly through flea bites.
   See Baldacchino et al. (2014).
   Diphtheria is an infection caused by the bacterium Corynebacterium diphtheriae. The German bacteriologist Friedrich August Johannes Löffler (1852-1915) was the first person to cultivate the bacterium.
   Reference to Otten (1936). Otten’s work with dead and live plague vaccine attracted much attention because it deals with the very fundamental question of the best and safest method of immunisation of human beings.
   Epidemic typhus is due to Rickettsia prowazeki, spread by body lice, scrub typhus is due to Orienta tsutsugamushi spread by chiggers and murine typhus is due to Rickettsia typhi spread by fleas.
   The reference is to the bacteriologist Louis Edmond den Dooren de Jong (1897-1972), den Dooren de Jong (1942).
   Mycetoma is a chronic infection of skin and subcutaneous tissue. The condition was first described in the mid-1800s and was initially named Madura foot, after the region of Madura in India where the disease was first identified.
   Possibly a reference to Cuscuta howelliana an abundant endemic parasitic plant that inhabits California vernal pools.
   Loranthaceae, commonly known as the showy mistletoe. Loranthaceae are primarily xylem parasites.
   Mistletoe is the common name for a number of parasitic plants within the Santatales.
   Feldmann not identified.
   Probably a reference to a publication of the Wilhelmshafen team of the Senckenberg am Meer Institute concerning the Wadden Sea tidal flat ecology.
   Cambial activity increases the girth of stems and roots by producing additional xylem and phloem.
   Baas Becking gives a personal version of the first part of Goethe’s Gesang Der Geister über den Wassern (1779). The original texts is as follows: Des Menschen Seele/Gleicht dem Wasser:/Vom Himmel kommt es,/Zum Himmel steigt es,/Und wieder nieder/Zur Erde muß es,/Ewig wechselnd.
   Baas Becking referred to Jossephus Jitta (1932).
   Baas Becking described the Carbon cycle in Chapter VI of Geobiologie (1934), (p. 61-64 English edition) and on p. 653-660 of the 1953 manuscript of Geobiology. About “Bacillus oligocabophilus” he remarked:
It is curious that Beijerinck was able to isolate an organism that can obtain its energy from the oxidation of CO to CO2 and that there is sufficient CO present in laboratory air to obtain good quality cultures of “Bacillus oligocarbophilus” on a substrate consisting of tap water to which a small amount of phosphoric acid has been added.
   Reference to Porter (1925). Charles Walter Porter (1880-1971, Professor of Chemistry University of California, Berkeley (1925-1946). Baas Becking probably was acquainted with Porter.
   Baas Becking described the nitrogen cycle in Chapter VI of Geobiologie (1934) (p. 64-65, English edition). In the 1953 manuscript of Geobiology Baas Becking described the nitrogen cycle on p. 668-672.
See for recent review of the Nitrogen Cycle (Thamdrup, 2012).
   See for Kingma Boltjes (1934) Section 4.7, for Martius and Knoop (1937) Section 7.1, and for Chibnall Section 3.9.1.
   On March 26, 1938 Baas Becking gave an address on the sulphur cycle during the fifth scientific meeting of the Dutch section of the International Society of Soil Scientists (Baas Becking, 1938b). See Baas Becking Geobiologie (1934), Chapter VI (p. 65-67 English edition, 2016); Baas Becking manuscript Geobiology (1953a, p. 661-668).
See also for the present state of the art of research on the Sulphur Cycle. Jørgensen (2021).
   Baas Becking used the word “philothion” in Figure 7.24 of the sulphur cycle. It referred to the finding of J. de Rey-Pailhade (1888, Comptes Rendus Acad. Sci. 106, 1683-1684) of a compound capable of producing hydrogen sulphide from sulphur powder. Nowadays known as reduced gluathion (GSH), present in most living cells from bacteria to mammals (except some bacteria and amoeba).
   Present name Acidithiobacillus thiooxidans
   Thiols are sometimes referred to as mercaptums.
   A reference to Henry James Bunker (1897-1975) and his review of sulphur bacteria in Bunker (1936).
Bunker’s specialty were the bacteria of the sulphur cycle, his research material being at first Thiobacilli and later the sulphate reducing bacteria. The sulphur bacteria were then a rather neglected group, studied mainly in Holland, and Bunker’s review collected together the scattered literature and laid the foundations for developments in this area after the Second World War. His review is perhaps the most recondite of Stationery Office publications to have achieved a second edition: it was reprinted in 1951. The sulphate reducing bacteria were then among the most awkward of the sulphur bacteria to handle, yet they showed the widest range of economic activities, one of the most dramatic of which was their role in the corrosion of buried pipes.
See Postgate (1976).
   Reference to Stephenson and Strickland (1931).
   Reference to Von Wolzogen Kühr (1923a, 1923b, 1937), Von Wolzogen Kühr and van der Vlugt (1951).
   Reference to Pauling (1939). See also Section 2.5.15.
   In the 1953 version of Geobiology Baas Becking remarked about the biocoenosis (p. 591):
A biocoenosis represents a polysymbiotic relation. Symbiotic relations are, to large extent, relations of ‘give and take.’ The great advance in the Neurospora research has done much to strengthen this belief. Loss mutations may account for a great many relations, but we should not forget that there are organisms which probably never possessed certain powers, necessary for their maintenance and their development and that these organisms, therefore, are “chronically” dependent on others.
On p. 635 of the 1953 Geobiology manuscript Baas Becking wrote in eloquent style:
If words are comparable to organisms, sentences should be like the biocoenoses, like living communities. We may also use another simile, that of music, where we have notes and musical phrases. Here the blending of the simultaneous and the successive is known in one form of music; the canon.
The biocoenosis, moreover, reminds us strongly of the ‘belles lettres’, for it is a story of mating, begetting, competitive maintenance, of dissipation and of farewell, both successive and simultaneous phases being represented. A biocoenosis of more than two components is already difficult to analyse. In still more complex cases we must often be content with descriptive or theoretical treatment. If we realise certain relations, we are prone to enlarge, and enlarge upon, their importance. If we cannot account for certain phenomena, we either allow them to stay unaccounted for (and many are the questions raised by our scientific ancestors which we treated with such neglect), or we may invoke “chance” (when mechanistically inclined) or “directed forces” (when vitalistically inclined) to take care of those, still indigestible, morsels. The great system of nomenclature, developed to describe the natural communities is very often but a cloak to hide our ignorance.
Metabiosis finds its highest expression in the biocoenosis, but metabiotic relations are of multiple nature and may be traced in many more elementary substrates.
   Jean Jules Beauverie (1874-1938), French botanist and mycologist. See also Section 7.1.
   Reference to Robert Warrington who presented early experiments with self supporting systems in aquaria in 1850 to the British Chemical Society: ‘On the adjustments of the relations between animal and vegetable kingdoms, by which the vital functions of both are permanently maintained.’ He placed two goldfish in 20 gallons of spring water, which half filled a glass bowl. Some sand, mud, pebbles and bits of limestone were added to the bottom of the bowl. The mud was used to hold the roots of Vallisneria. Warrington left the aquarium undisturbed until a green scum started to coat the wall of the bowl, obscuring the view of the goldfish. The addition of some snails, which fed on the green scum and decaying vegetable matter resulted in clearing the water.
   Reference to Justus von Liebig (1803-1873), German chemist. Liebig demonstrated during popular lectures in the Aula of the Munich University a Warrington closed aquarium system. Liebig had set up an aquarium for Queen Maria von Bayern (1811-1864), a “Liebigsche Kleine Welt im Glase” for her Salon in the Munich Royal Palace.
   Hindu Gods: Shiva (destroyer) and Vishnu (protector).
   The Amsterdam Zoo ‘Artis’ has a seawater aquarium since 1882. Unique to the ARTIS Aquarium is the William Alfred Lloyd water filter system that is still working today. With its masonry and tube labyrinth, this system has been supplying the entire Aquarium with naturally filtered water for almost 140 years, which makes fresh and saltwater life possible in the more than 60 different aquariums.
In Geobiologie (1934, p. 176), Baas Becking referred to the Amsterdam PhD thesis of Catharina Honing (1933), Onderzoek over de Reiniging van Zeewater in Groote Aquaria.
   Present name: Desulfovibrio desulfuricans.
   In Section 6 Baas Becking referred to Hecht (1933). Reference to Correns (1939, p. 209-210).
   Baas Becking referred not to Dorothy Stephenson but to Majory Stephenson (1885-1948), British biochemist, who wrote the classical textbook Bacterial Metabolism in 1930. Baas Becking referred to Stephenson and Strickland (1931). See also Sections 4.8 and 7.8.8.
   See for reference to research of Wolzogen Kühr, Section 7.8.8.
   Reference to Verhoop (1940). See also Section 6.
In the 1953 manuscript of Baas Becking remarked (p. 666-667):
Verhoop (1940) gives a colour scale of various blackness, corresponding to natural muds. The “palest” mud, where the FeS may just be discerned, corresponds of 0.1 mg FeS per kg soil mud, where the FeS may just be discerned, corresponds of 0.1 mg FeS per kg soil, 18 % moisture. This would mean about 0.1 γ per cc. The blackest mud corresponded to 320γ/cc sulphur. The values assumed for the “terrestrial” intensity of the sulphate reduction are of the same order of magnitude. In oxidation of this reduces sulphur, 4 x 108 tons of oxygen would be fixed. Inasmuch as the total oxygen in the atmosphere amounts to 1.2 x 1015 tons, the sulphate reduction cannot, except locally influence the oxygen tension on a large scale.
   ‘Walvis Bay’ in Namibia, a safe haven for sea vessels, rich in plankton and marine life, attracting large numbers of southern right whales. Ruppia maritima is an estuarian pondweed in Walfish Bay. Baas Becking possibly referred to Brongersma-Sanders (1943).
   Oren (2011, p. 21-22), discussed Baas Becking’s fascination for the brine shrimp Artemia.
   In the 1953 manuscript of Geobiology Baas Becking referred to the Boekelo research (Nicolai and Baas Becking, 1935) (p. 136-137):
There are rare as well as common microbes. But usually the forms will fill their “niche” in a surprising short time. A case in point is the salt bath of Boekelo, in Holland. The salt in Boekelo is obtained by pumping water into the subterranean layers of salt and pumping up the brine. In 1934, exploitation of the pool was started, after infection with seawater. In August of that year the author could find, however, no specific marine organisms in the pool. A few microbes typical of strongly saline solutions were present. In the last days in August 1934 a mass development occurred of a rotifer, a Brachionus, approaching closely in form the Brachionus described from the salt lakes in Siebenbürgen and found by myself in Soda Lake, Nevada, 1924.
Soda Lake was conspicuous for the mass development of brine flies (Ephydra sp) the pupae of which formed veritable floating islands (they are roasted and eaten by the Indians). While this fly did not occur at Boekelo in 1934, it suddenly made its appearance, in such masses that it hampered the use of the bath. The closest “saline” stations were; Greifswald (Pomerania, near the Baltic), Mulhouse (Lorraine) and Staffordshire (England). It seems most probable that a complete biocoenosis, corresponding to the chemical composition of the milieu, would have been completed in due time.
See for Boekelo also Section 4.3.9.
   In the 1953 version of Geobiology Baas Becking remarked (p. 298):
In the sediments the lake writes down its history. In desert lakes as well as in glacial lakes sediments are often “varvate”, i.e. laminated and stratified. If the annual season contribution be known, measurements on sediment will give a clue to the previous history of the lake. Important “markers” are the aeolic pollen grains. Their nature and frequency in a so called “pollen diagram” may yield important clues as to the age of the deposit. This is particularly important in lakes of glacial origin as the Wisconsin Lakes and the Connecticut Lake ‘Linsley Pond.’
   See for Zürich See and Minder, Sections 5.9 and 8.5.5. For Walfish Bay see Brongersma-Sanders (1943) and Section 5.12.3.
‘N.W. Polder’ [Noordwestpolder]; initially the name for the present Wieringermeerpolder, newly created land, developed in the 20th century by draining parts of the inland Zuiderzee. Draining was completed in August 1930. After desalinisation the new land became usable in 1934. See Baas Becking (1936a, 1936b).
   See Sections 7.9.1 and 7.9.2.