5. INFLUENCE OF THE MILIEU UPON THE ORGANISMS

The living mass of one organism, or a population seems to increase according to a special law. The increase is often first fast, then slows down, until growth stops. There seems to be for the organism and for the population a limit which is slowly reached. Robertson compared the process to an autocatalytic reaction in which the reaction velocity dy/dt = y (a-y) in which y is the amount of substance transformed, and a the limiting amount of substance. Now in the case of organisms there is such a limit (obviously). It is not said, however, that organismal growth is a reaction with a single “master-catalyser.” For population growth, however, the imputed limit seems very questionable. There should be an “overhanging” fate over a population, a sort of doom, represented in the a of the above differential equation. R. Pearl (1926) actually seems to suggest such a thing.¹

The equation of the hyperbolical tangent is (Kavanagh and Richards, 1934),

really the general expression of the “autocatalytic” population curve of T.B. Robertson and of Raymond Pearl. For if we take a population curve:

and we write:

shifting the coordinates, we obtain, for the right hand expression:

For the Robertson curve:

we determine the first derivative as

the second derivative gives:

the third derivative:

Now the first derivative represents an optimum curve, maximum at z = 0, points of inflection at the values for x in:

or,

[Baas Becking inserted Fig. 5.1. with the Kavanagh and Robertson and Pearl sigmoid growth curves.]

In his book, The Chemistry of Large Molecules (1939) Mark gives examples of the growth of polymers.³ This is a case from inanimate nature much closer allied to our problem as catalyst reactions. Mark finds too a few cases that this growth may be represented by F-shaped curves. In the majority of cases hollow, or linear relations appeared. It follows that already here conditions are very complex and elude analysis. Therefore, in growth, where conditions are so much more involved, it is very risky, like Pearl does, to dictate a certain shape to a curve which obviously does not possess it!

Now there are a great many similar curves. The integral of the normal curve

is an example, the integral of:

and a great many other expressions the treatment of which here would require too much space. The author tried the goodness of fit of various population curves with twelve expressions and found only in a few cases a tolerable fit for the autocatalytic curve. This shows that the S-curve, being of very common occurrence, may have a variety of causes. This is also demonstrated by the fact that several distributions seem to fit the ascending branch, or the inverted descending branch of the normal curve. There is one case, however, where cause and effect has been studied (Baas Becking and Baker, 1926) and where the ogive, or rather the point binominal fits the observations most closely.⁴

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The laboratory milieu does not even distantly approach to the potential milieu, for the simple reason that it is impossible to delineate the potential milieu. The polyblepharid alga Dunaliella is able to live in 1 molar solution KCN. I only tried a limited number of compounds on Dunaliella and in how many organisms we tried the effect of a solution of 1 molar KCN? Therefore, we would rather be silent for want of facts if not, even at this stage of our science we might perhaps make certain generalisations. When we for example decimate the milieu of the filamentous alga Chaetomorpha linum in mixtures of 3 cations Ca, Mg, Na and three anions HCO₃, SO₄, Cl, we find its milieu optimum at a point which is not represented by any “terrestrial solution.” The case seems to suggest a similar conclusion as the weird liking of Dunaliella for prussic acid. The relation of the earth, and its present geochemical condition and the organism is much lower than we had expected.

The chemical and thermal milieu are dependent upon water. For the chemical milieu “corpora non agunt nisi solute” hold,⁵ and we have seen that thermal action on dehydrated cells is materially lessened. Therefore, the condition of water in the cell seems again to dictate. If a spore or a cyst were perfectly dry, perhaps it could start on an interplanetary trip, carried by radiation – pressure – as Svante Arhenius assumed. But radiation remains as an important agent. It remains to be seen whether the enormous intensity of ultraviolet, lethal radiation, effectively screened off by our atmosphere, would not kill anything, however dehydrated, that came under its influence! Henderson has called special attention to the thermal properties of water which properties caused this earth to process its lasting and equable climatic conditions. The gravitational field of the earth, finally determines the composition of atmosphere and hydrosphere. As a matter of fact, for our living beings in the “hydrosystem” (there is, as we have seen, also an ammono system), the earth is a pretty fine abode. But we should not forget that we are here because the conditions are so favourable, and that these conditions were not created to pleasure! (Whewell).⁶

Stenobionts are very fragile organisms. They live within very narrow milieu boundaries. They live, to use the words of Robert Bridges (Testament of Beauty) “on the sharp of a razor, that may not e’en be blunted, lest they sicken and die.”⁷ Of a great many organisms, which we have only observed, but never experimented with, we suspect this stenobiotic nature. There are, for instance, mosquito larvae occurring in the liquid of the beaker of the insectivorous plant Nepenthes (Tropische Natuur, 1929).⁸ We do not know whether there are stenobionts, only experiments will show. In relation to temperature J. Ruinen has investigated the green Ulothichal alga Ctenocladus circinnatus.⁹ This alga is able to develop in a temperature range of 16-21 °C. This is a very narrow limit.

The tunny [also called ‘tuna’] is a fish that cannot withstand temperatures lower than 14 °C. Many instances of flowering are to be ascribed to temperature influence in a previous developmental stage. Jarovisation (= vernalisation) of wheat is one of the examples here.¹⁰ Many Desmidiaceae have a relation to the pH, occurring only in acid waters. For animals the relation to acidity is not so pronounced. The alga Lochmiopsis (Ctenocladus) mentioned also has a very remarkable saline milieu, being extremely calciphobic, alcalophilic and osmophilic up to 1 molar NaCl, beyond which the organism cannot exist. Its temperature milieu is even more limited akinetes germinate only between 16-20 °C.¹¹ Several organisms are closely fixed to the osmotic value of the marine environment, although there is more surmise and talk about this than experiment. The marine plankton crustacea Evadne, for instance, lives for days in distilled water!¹² The most beautiful examples of stenobionts one finds in parasites. Even temporary ectoparasites like body lice, show already, a great conservation and apparently, have never left their host for millions of years (Ferris, De Anoplura).¹³ How else to account for the fact that the louse of the camel and that of the llama are closely related? (see also Section 7). Stenobiontic life is therefore often closely related to dependent life, and the closer the dependence of an organism upon another, how narrower the milieu becomes.

[In the margin: A typical stenobiont is the alga Hydrurus foetidus, the water tail, only occurs in water with very little electrolyte. This as opposite to halophytic organisms which are, however often eurybionts (Dunaliella).]

A real eurybiont one should not miss in extreme conditions of the milieu. Whether acid or alkaline, cold or hot, freshwater or concentrated brine, the organism should be there. When the earth should be heated up or cooled down, the eurybiont should keep the earth company to the bitter end. Let us see which organisms we find.

a.Bacteria.Sulphate reducing bacteria I found in a heather bog, pH 3.8, also in Searles’ Lake, Nevada, pH 10.8. Under the ice we find sulphate reduction but also in the hot springs at Kali Pait (48 °C).¹⁴ The bog water contained almost no minerals, while the solution at Searles’ Lake was saturated. Cellulose bacteria and biologic acid Closteridia probably also fall under the eurybionts.

b.Protozoa.Amoeba we find everywhere, in very clear mountain water and in concentrated brine (Baas Becking and Ruinen) in Lake Tyrrell, S. Australia pH 8.8 and in Soda Lake Nevada pH 10.8.¹⁵ In the hot springs of Kawah Tjiwedéh (Java) and in Mammoth Hot Springs, Yellowstone up to 58 °C. But also, in the cold ditch water collected over ice in January.

c. Bluegreen algae. Accompanying everywhere. Hof (1935) described them from brine and Beijerinck from bog water (1902).¹⁶ They also occur in a pH range 3-11. In hot springs of Yellowstone van Niel found these up to 75 °C. In the lichens they occur in temperature of -40 °C.

d. Nematods. Apparently have a very wide milieu. There has hardly been any large sample of extreme milieu that did not yield one or more of these curious animals. They are worth investigation, especially those living in hot springs and in saturated brine (Australia, Lake Bumbunga).¹⁷

e. Fishes. It seems that, apart from certain flies that could be mentioned here, fishes are apt to withstand extreme milieu conditions. Gasterosteus lives in acid bog water without minerals, but also in an alkaline 10% NaCl solution. Cottidae live, according to Hecht, in a seawater entirely deprived of oxygen, and also in hot springs. Cottidae in Alaska are reported to have withstood solid freezing at -40 °C for a winter.

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The fundamental law of photochemistry (Grothius-Draper) states that light, in order to act, should be absorbed. This means that the living state should show absorption bands at the places of the spectrum where radiation is utilised. Another fundamental law of photochemistry is given by Einstein, where the energy necessary for a photochemical reaction may be expressed as 𝛥t = h𝜈1 – h𝜈2 if the frequency 𝜈2 be 𝜈c radial (fluorescence) 𝛥E be positive 𝜈1 should be >𝜈2 from which follows that the wavelength of the fluorescent light should be larger than that of the absorbed light (Stokes’ law).

[Baas Becking inserted Fig. 5.2.]

Bluegreens, purples under quartz, under salt.¹⁸

[Baas Becking inserted Fig. 5.3, a small drawing of light spectra green and purple bacteria.]

Eye of Gyrinus.¹⁹ Melanophore contraction.

[Baas Becking inserted a rough sketch of light spectrum of the eye of Gyrinus (Fig. 5.4).]

Light and shadow lenses. Etiolation.²⁰ Other influences.

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[Baas Becking inserted a rough sketch Fig. 5.5.]

[Baas Becking inserted Fig. 5.6, light spectrum in relation to sensitivity of organisms.]

Temperature is nothing but the statistical average of the velocity of the molecules in a system. The absolute void has no temperature and interplanetary space has only temperature in so far as it can burst on material particles. Organisms are excited by, or put to sleep by temperature, not merely because they consist chiefly of water and the properties of water change so much with temperature, but because equilibria are upset, (sweetening of potatoes, work of B.D.J. Meeuwisse).²⁴ Only a long monograph would do justice to this subject matter. In this section we shall only select a few high spots and deal with those, of course rather superficially. But the argument should not be too long.

A great many organisms, especially in spore form, are able to withstand extremely low temperatures (see Baas Becking, Geobiologie, 1934). Vital functions, however, should stop whenever the liquid phase disappears. D’Hérelle (oral communication) found an Aspergillus in a brine bath at -35 °C. Systematic study of the behaviour of organisms at very low temperatures seems not to have been carried out. Necrobiotic effects take place at -40 °C, while in deep freezing fruits it may be observed that the oxidase activity (Cu-containing proteid; Kubowitz)²⁵ the peroxidase activity (Fe-containing porphyrine proteid, Kylin)²⁶ and the katalase activity not only remain unimpaired, but continue, very slowly, at these improbable temperatures.²⁷ Also, amylase activity could be demonstrated. It has been suggested (Gortner,²⁸ Kruyt, Baas Becking) that part of the water remains in the non-frozen condition at very low temperatures (far below the freezing point). To call this water “bound water” makes more claim for our knowledge of, and insight into the effect, than we desire (Baas Becking, 1942b).²⁹

[Baas Becking inserted Fig. 5.7, the link between intensity of a biological phenomenon, time and temperature.]

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[Baas Becking inserted Figs. 5.8a and 5.8b (both schemes from Shelford, 1929).]³⁰

Dry interior U.S. (1900-1912). 71,000 deaths (Huntington).

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Natural milieu contains only a few acids and bases which exert a dictating influence, although, actually the number of acids and bases in natural environment is legion.

Inorganic acids

1. H₂CO₃, 2. H₂S, 3. HNO₃, 4. H₂SO₄, 5. H₃PO₄, 6. HCl.

Inorganic bases

1. Na₂SiO₃, 2. Al(OH)₃, 3. NaHCO₃, 4. Na₂CO₃, 5. Na₂S, 6. NH₃.

Of the organic substances we shall deal only with five

1. Oxalic acid, 2. Humic acid, 3. Butyric acid, 4. Lactic acid, 5. Acetic acid.

HCO₃ is, of course, the ‘photosynthetic acid.’

Silica may be toxic to certain organisms. Apart from human siliciosis. Dr. P. Schure found glan, mica and even quartz toxic to swarmers and myxamoeba of Reticularia lycoperdon.³¹

Formation [of oxalic acid] obscure. Breakdown by a curious enzyme (Fig. 5.9). Zaleski isolated the enzyme from wheat. Kruyt says plentiful in mosses. Bassalik bacteria.³² Franke and Hasse more recent study. Niekerk (unpublished).³³

and with access of water

and with access of oxygen

It is very remarkable that H₂O₂ should be formed here, while in mosses there is catalase present. The enzyme is highly thermostable and works in highly acid media. Bacteria extorquence in the gut of earthworms (Bassalik, 1913) is also able to attack Ca(COO)₂, one of the stable ‘organic compounds’ (only soluble in pH < 2. It is therefore probably the only organic acid that enters in appreciable quantities in the carbon cycle (leafy humus)

May be formed from CO and H₂O₂? Better a scheme of Chibnall (1939).

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After the classical work of de Vries (1883-1884) and Hamburger (1884), the osmotic value of the milieu, whether hypotonic or hypertonic in relation to the inner environment, has been recognised as a factor of prime importance.³⁵ Marine animals brought into freshwater take up an inordinate amount of water, which they have to recrete. This fact, originally considered as a purely osmotic phenomenon, requires other concepts for its final understanding (see Section 4.2.5.a). Also, the fact that organisms may exist in saturated salt solutions, and still are able to take up water, cannot be accounted for by osmotic theory. The phenomenon of swelling and of antagonism have to be taken into account to create a harmonious theory of water intake and of water recretion. It should never be forgotten that the classical osmotic theory only refers to water movement and that all other phenomena require theories on intrability and on permeability. It may be stated in advance that the pressure of at least two semipermeable membranes in a plant cell has been proved.³⁶

Van ‘t Hoff, using the data of W. Pfeffer (1877) established the fact that the osmotic pressure, expended [exerted] by a sucrose solution against a semipermeable membrane (Cu, ferrocyanide), followed the gas laws (Fig. 5.10).³⁷ PV = RT or rather, per unit volume P = cT, in which c is concentration. A one molar solution of cane sugar may generate a pressure of 22.4 atmospheres. De Vries found, by means of his “plasmolytic method” that electrolytes acted as if they were much more concentrated. The “isotonic coefficient”, a multiplication factor to satisfy theory, proved to be, for NaCl almost 2, for Ca(NO₃)₂ almost 3 etc. Arrhenius (1887) from these data and other cryoscopic and conductivity measurements, derived his ionic theory. The ions behaving as separate particles in the solution. A saturated NaCl solution, practically 100 % dissociated and approx. 5.25 molar, should yield an osmotic pressure of 10.5 × 22.4 = 235 atmospheres.

(Dubois, Herrera, Traube).³⁸ The copper ferrocyanide membrane, but particularly the metal-silicate membrane is somewhat plastic a crystal of nickel sulphate in a solution of sodium silicate will form a semipermeable membrane of nickel silicate, surrounding the crystal. The high concentration of Ni²⁺ and SO₄^2- ions within the membrane will cause a flow of liquid towards the crystal. The membrane is stretched, bursts and new membranes are formed. In this way curious plant-shaped structures may be formed. However, it is as futile to compare these structures with vital growths, as it is to account for the annual rings in the trees as a Liesegang phenomenon.³⁹ A plant cell will take in water till the turgor equalises the tension of the cell wall. Growth proceeds when, by means of growth hormone, the plasticity of this cell wall is increased, and the turgor pressure may again become active in stretching the plastic wall till it reacts again to the renewed tension.

Several protozoa, myxomycetes and conjugate algae possess a pulsating vacuole which recretes water at a certain rhythm (Fig. 5.11 after C.V. Taylor (1923), on Euplotes).⁴⁰ In a freshwater milieu the rhythm is frequent, transference in a milieu of high osmotic value causes the frequency to decrease (data from Pelseneer, and Yves Delage).⁴¹

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Published in the Arch. f. Mikro[biologie].

Noel Patten, Nereis,⁴² LBB Evadne.⁴³

M. Fischer.⁴⁴

Water intake cannot proceed by osmotic forces only. The swelling of biocolloids has to be taken into account as well.

At the outset it is well to point out, that the word “antagonism” is used by pathologists in a different sense, viz., the inimical action of certain microbes towards one another. The word however, is pre-empted by physiologists to indicate originally the phenomenon that 2 substances, A and B, in themselves toxic, may appear to exert a beneficial influence upon an organism when combined (Fig. 5.12). Now pharmacologically the effect may be, moreover additive, as the two toxic agents may, in combination, enhance each other’s effect. In the latter case we meet with sensitisation. There are many in analogues in physical chemistry. Boiling point, melting point, viscosity, etc., if a brisant mixture may be either additive or non-additive. This is particularly striking in the case of alloys. Physiological antagonism is important to geobiology as the oecomena of organisms in natural mineral milieu depend, for a large part upon the antagonistic action of the combined mineral components which may, in themselves, be highly toxic. In the milieu interne we meet, moreover, with body fluids which show similar properties. It will be seen that antagonistic rather than osmotic effects dictate the boundaries of the milieu externe for a great many organisms. Although the study of the concept enjoyed its highlight in the school of Jacques Loeb, a renewal of interest would be very expedient, as the great number of most striking and suggestive observations have been made, which still await a basic, comprehensive, theory to account for them.

Sydney Ringer, in 1881, succeeded in keeping a frog’s heart beating by perfusion with an extract of dried ox blood.⁴⁵ Later he used a weak solution of common salt. Repeating the experiment with a solution of NaCl in distilled water, the heart stopped beating. It is Ringer’s great merit to have recognised the nature of the phenomenon: NaCl in itself was toxic, but it became detoxified by the calcium in the London tapwater. Ringer also recognised the two types of solution needed; 0.9 % for warm blooded, 0.7 % for cold blooded animals, and the relation of the Na:Ca, in atmos 40:1.

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The blood of the mammal is an archaic seawater, taken from the primordial ocean when in evolution, the aquatic animal became a land animal. Now it is true that the freezing point depression of the blood of cartilaginous fishes (rays, sharks) varies but little from that of the surrounding seawater, unfortunately that serum is no seawater as the osmotic regulation is performed by means of urea.

[Fig. 5.13 see also Figure IX.6 in Geobiology, 2016.]

Seawater [corrected in purple pen by “blood”] has proportionally much more Mg and HCO₃ than seawater. It looks like a seawater which has exchanged bases with soil (Rockanje Lake, Island of Voorne, oil waters of Tjepin and other subsoil wells, central Java).⁴⁸

If this serum has anything to do with a natural water it is certainly not seawater. Now it may be claimed that “while the ocean got ‘saltier and saltier’ the ionic points travelled in the triangle” as indicated in Figure 5.14 for a trip down river towards the ocean. This is an unsafe assumption, as the NaCl in the ocean cannot be accumulated there by weathering alone (see Sections 2.3.1 and 2.4.6). We do not know how the ocean came to be. Probably volcanic action extends into the game at certain instances. And taking the concentration of the blood in NaCl 0.62 %, the triple [?] point of such a natural water would still be situated right near that of seawater (see Section 5.7.4 and Fig. 5.14 for freshening of the Zuyderzee). The hypothesis therefore is clever, but exceedingly improbable.⁴⁹

Jacques Loeb claimed that, seawater was, as far as ionic balance was concerned, “an ideal abode.” Osterhout went so far as to culture wheat in diluted seawater. What probably happens in seawater is that it behaves like an “antagonistic buffer”, dilution and concentration over a small sample did not influence the ionic balance. But any other claim as to the superiority of seawater no experiment has, as far as the author knows, substantiated.⁵⁰

The composition of seawater is dictated by organisms (Fig. 5.14 and Table 5.2). [The figure shows the changes in percentual composition of average river water R to oceanic water, for 8 components.]

• Silica disappears, bicarbonate almost disappeared

• Sulphate decreases

• Calcium decreases

• C = Caspian Sea (diluted with Wolga and Ural water)

See also Figure 6.3 and Section 6.4.2.c.

• 1) The biologically active ions have all decreased on their journey

• 2) Na and Cl, biologically much less active, have accumulated

• a) HCO₃ ion. Decrease due to photosynthetic active lime precipitation and oversaturation.

• b) SiO₃. Probably entirely removed by organic agencies.

• c) See HCO₃.

• d) SO₄. Sulphate reduction!

• e) K decrease still more.

According to Clarke (1916) very active, p. 146, “the biochemistry of the ocean is curiously complex, and its processes are conducted on an enormous scale.” 300 million tons of sulphate being precipitated (reduced!). Surface earth ≏ 5.12 × 10¹⁸ cm². Complete reduction 96 g SO₄ yields 64 g oxygen or 200 × 10⁶ ton. This would make no material difference in O consumption (see Section 6.4.3). Also see Figure 3.24, where more material has been brought together.

[Baas Becking inserted Figs. 5.15 and 5.16.]

Literature

• Baas Becking and Boone (1929) [= 1931].⁵¹

• Baas Becking and Jacobi (1931) [= 1933].⁵²

• Baas Becking, Karstens and Kanner (1934) [= 1936].⁵³

• Booij and Bungenberg de Jong (1934) [= Bungenberg de Jong, Booij and Wakkie (1936)].

[The triangular diagrams are useful to illustrate boundaries of ionic milieu. Here the ionic antagonism plays a role, the cations being the most active in this respect. If total normality of the chloride constant (= 1 in the Fig. 5.15) the shaded area approximately represents the area in which eggs of Artemia salina, the brine hatch. This area is different for various normalities.

If concentration is used as a vertical axis we obtain a triangular prism, the cross sections of which are equilateral triangles, each corresponding to a certain normality (Fig. 5.16). For the hatching of the brine-shrimp eggs, between normality 0-4 we obtain the following figure, the solid, shaped like a part of an orange corresponding to the salt combinations in which hatching of the eggs is possible. It has been shown by Bungenberg de Jong and his coworkers that suspensions of lecitine assume a negative charge in certain three chloride combinations, while in other combinations the charge is positive, the ‘negative’ area coinciding, more or less, at 1 normal, with the area in which the crustacean eggs germinate.]⁵⁴

[Baas Becking inserted Fig. 5.17. in which he indicated the potential environment for three chlorides for the fungus Oöspora. See also Figure X.18 in Geobiologie edition (2016). See also Figs. 5.18, 5.19b and 5.20.]

T. Hof, PhD Dissertation, Leiden (1935).⁵⁵

Curling of liana[?].

J. Ruinen, PhD Dissertation, Leiden (1933).

[Baas Becking inserted Fig. 5.18, potential environment for three chlorides for Lochmiopsis sibirca, based on Figure X.18 in Geobiologie (2016). See also Figs. 5.17, 5.19b and 5.20.]

Baas Becking (1930, 1931a and 1931c).⁵⁶

[Baas Becking inserted Fig. 5.19a and 5.19b, Antagonism of Dunaliella between calcium and magnesium with evaporation of seawater, based on Figure X.20 in the 2016 edition of Geobiologie. In Fig. 5.19b he also indicated the potential environment for Dunaliella for three chlorides. See also Figs. 5.17, 5.18 and 5.20.]

Chaetomorphalinum (J. de Zeeuw, PhD Dissertation, Leiden, 1937).⁵⁷

[Baas Becking inserted Fig. 5.20, in which he indicated the potential environment of Chaetomorpha linum and other algae salt organisms for three chlorides. See also Figs. 5.17, 5.18, 5.19a, and 5.20.]

The unpublished work of Reitsma and also the thesis of H. Booij have shown that the pollen of sweet peas, while readily germinating in fine cane sugar solution is greatly stimulated by CaCl₂ in concentrations up to [50 m.eq].⁵⁸

[PhD Dissertation, Booij (1940)].

[Baas Becking inserted Fig. 5.21, based on the dissertation of H.L. Booij, in which the potential environment was indicated for the germination of Lathyrus pollen for a mixture of three chloride solutions. See also Figs. 5.17, 5.18 and 5.19a.]

Theunissen.⁵⁹

[Baas Becking inserted Fig. 5.22, a rough sketch that is a summary of the foregoing paragraphs on the potential environment of various organisms in mixtures of chlorides. The sketch was copied from Figure X.18 in Geobiologie (2016).]

If the milieu should act directly on the genome, we might expect a world extremely Lamarckian. The imprint of the milieu would result in a hereditary effect. Although many Latin biologists adhere, in some modification or other, to Lamarck, experimental evidence, however, has in no case substantiated this theory. (Claims to a Lamarckian origin of hereditary change, such as from Guyer⁶⁰ and Kammerer⁶¹ have been disproved). We find ourselves, therefore more or less in a quandary. There is no doubt that organisms fit into their milieu, a survival of the fittest cannot account for the presence of highly specialised organisms in a highly specialised milieu, as the probability to live is so infinitely small as compared with the probability to miss. Still all attempts to demonstrate a direct action of the milieu upon the genome have failed. There is, probably, an indirect action (indirect in as much it bears no relation to the fitness of the new genome to the milieu) of both cosmic rays and certain chemical substances of the colchicine or acetophenone type. Cosmic rays seem to promote mutations, at least in Drosophila, which the colchicine-like substances may cause polyploid mutants to occur (Krythe and Wellensiek, 1943).⁶² By examining the actual nature, however, we are struck by the remarkable fit of external and internal milieu, a fit that none of the theories mentioned may explain. Ignorabimus!⁶³

This word is the bugbear of many biologists, and the concept of teleology has indeed been treated often in an unscientific way, implying a function for every conservable biological structure. In this direction the inspiring book of Haberlandt, Physiologische Pflanzenanatomie,⁶⁴ has gone perhaps a bit too far, to be silent about many investigations on flower biology. However, living beings behave not aimlessly, they apparently strive for some goal. The goal, the end the aim, lies primarily in themselves, as already Aristotle recognised. The concept of entelechy (H. Driesch),⁶⁵ should be taken more seriously than most workers seem to do, in as much as it is really capable to stimulate further research. For what is “vital pressure”, élan vital, but our expression of our entelectric principle? From this tendency for self preservation (self used here as the species rather than the individual) we arrive automatically to teleological ideas. Adaptation is a change, a useful change, therefore teleological, of an organism. It is a change caused by the milieu. It seems as if such a change is often hereditary and then we are up head and shoulders into Lamarckians. For it is different if a variation range of a given organism is large enough to select from (Darwin) or whether as a given pattern, a new pattern is superimposed.⁶⁶

The word “physiological artefact” was coined by A.J. Kluyver and J.K. Baars, to describe sulphate reducing bacteria, isolated from natural surroundings, which proved to be thermophilic.⁶⁷ Did the physiological experiment add something intrinsically new? Did the microbe mutate, or was there first a selection from a mass of variants, some of which die off because their potential milieu did not cover the conditions? We cannot strictly speak about mutation in microbes, for there is no sexual reproduction. Vaas has coined the word hypartype corresponding to phenotype and doxatype corresponding to genotype, instead of mutation he uses the word pedema.⁶⁸ According to Kluyver we have a cell A, potentialities a, changing into A, potentialities b. According to the statistical theory cell A would give rise to cells of potentialities a - - - z, out of which only a few meet their proper milieu. It is the process of adaptation that is difficult to understand in Kluyver’s theory.⁶⁹

Vaas (1938) has studied the effect of variable salt concentrations upon the growth of Bacillus megatherium de Bary (Fig. 5.23). This spore-former is a large rod, often in clusters or strings. The growth curve from a one spore isolation was in all cases investigated. Starting with a certain inoculum first a decrease in numbers (established nephelometrically) could be found. Later we see the so called “logarithmic phase” (Rahn, Buchanam and Fulmer) of growth, then a slackening off, a decrease in numbers followed by weak secondary maxima (line a). Line b represents the behaviour when inoculated in 9 % NaCl.⁷⁰ The initial decrease is much more marked, and subsequent growth is slower. By ingenious check experiments Vaas (1938) arrived at the conclusion that the probability for the occurrence of halo-tolerant variants depends upon the size of the inoculum. This he interpreted by the plausible assumption that variants of extreme potential milieu occur more rarely than variants of more average potentialities.

[Baas Becking inserted Fig. 5.24.]

At 0, on the abscissa the normal milieu, the variants able to withstand + or – might decrease according to probability law:

Now we ask for the potentiality +a it is clear that a population I is too small to yield such a variant. We need the sign of population II. From this Vaas (1938) concluded that these are no physiological artifacts, but only selection of extreme variants. (See however, the work of T. Hof, thesis, Leyden, 1932 on the halotolerance of urea bacteria).⁷¹

Heredity enters in here and with heredity comes phylogenetic and ontogenetic speculation. We meet with Darwinism. When selection is made by the milieu from a number of non-directed variants (as in the case of the halophilic bacteria), we meet with Lamarckism, where the milieu causes a directed change in the genome. Further we meet with the hybridisation theory of Lotsy and with mutations.⁷² Hybridisation may increase vigour as well as range of variability, but does not introduce the elements expressing the directive influences of the environment. So, speak about mutation only to refer to a sudden change in the genome, usually a loss mutation, while most adaptations, teleologically and biologically, are anything but loss mutations. By what curious process did the organism fit into its environment? Certainly, there must be discovered a new principle, for what we have does not suffice. Why is the eye of the water beetle Gyrinus partly adapted to land and partly to aquatic life?⁷³ How may we possibly account for such things?

This expression was coined by Troll (?) for, I believe, the relation between gall formation and the gall insect.⁷⁴ In gall formation we meet with a typical “ergon” action of the gall insect. In certain cases (Annand and Baas Becking, Science, 1925) galls have been produced artificially (cabbage leaves, ammonia vapour).⁷⁵ However, the insect must secrete the specific ergon to stimulate the plant tissue to make the gall. For a detailed description of the gall formation through the agency of Cynips, the classical memoir of Beijerinck should be consulted.⁷⁶ Zweckmässigkeit [Expediency] implies teleology, but what of that? The plant reacts in a way such as to further the ends of the animal, and without its reaction to the injection the insect could not produce larvae from its eggs. The insect is “adapted” to the plant, the plant to the insect.

Admission of the existence of mimicry, apart from a negative “selection of the unfit” of the unprotected, is tantamount to an admission of ignorance as to the cause of an orchid flower is able to imitate a female digger wasp, inclusive of the specific odour (which odour attracts the male digger wasp). It seems a phenomenon so far removed from any attempt at rational explanation, that we are reduced to a state of more or less dumb admiration. This is, perhaps, an unscientific point of view to take, but it seems to the author that the safe, so called scientific view is a hothouse flower, a product of the laboratory, where we select our problems arbitrarily and shrew the complicated ones.

A photographic picture; no – more than a photographic – a three dimensional picture of one organism has to leave its literal imprint upon the genome of another, so that the other shall wear the livery of the first. This as far as the efficient cause, and not the final, is concerned. The final cause is much more the component – mimicry is camouflage, an attempt to pass for somebody else. The final cause is, however, also, far more obscure. For who or what not only perceives the likeness between A and B, but is also able to create likeness and unlikeness?

Is not Francis Bacon right when he proclaimed final causes, like vestal virgins, as sterile and dedicated to God? For sterile is all our speculation upon these, and analogous matters. But certainly, it also points to an origin as far distant from our modern knowledge as the genetic relation is between the birch moth and the birch, or the Sphinx and the honeysuckle. In a sense these phenomena only allow for a transcendental explanation, as they transcend our understanding. All experiments upon the formative influence of the milieu have failed. Kammerer’s claim, that salamander (Axelotls?) raised in a dark aquarium and consequently dark skinned, produced dark skinned progeny, was based upon a falsification.⁷⁷ We stand utterly helpless to account for the simplest instances of obvious adaption. And we may not look the other side, for the phenomenon persist? And they belong to the realm of biology?

Geochemically it seems rather arbitrary to talk about minimum (or trace) elements. For the frequency of many of the ‘common’ biological elements (like carbon) is geochemically speaking, quite low, while geochemically common elements (nickel, titanium) appear as biologically rarities.

In this section we shall only deal with the beneficial effects and with the accumulation of the following elements:

Li, B, F, Ti, V, Mn, Fe, Co, Ni, Cu, Zn, As, Br, Mo, Sb and I.

There are references to other elements as well in the vast and rather secluded literature on the subject (Edelman, Goldschmidt). An element may be called “rare” when its dissipations lag is light (dispersion of Edelman).⁷⁸ This dissipation is again determined by its process of isomorphic replacement (Goldschmidt, 1923), meaning that a certain element may replace another, commoner, in its space lattice, if the ionic radii are of almost the same size. Due to this fact iron, which is really a central terrestrial element, ‘creeps up’ into the sima as its ionic radius is so close to that of magnesium. Due to its isomorphic displacement, again manganese replaces the iron in minerals. Many rare elements obtain, by this process a wide, but diffuse, distribution.

Often, the rare elements act in very low concentrations and are not further concentrated by the organism. Here is therefore a great difference between “éléments concentrateurs” (Vernadsky) and “rare elements.” In several desert plants, e.g., Astragalus, selenium, may accumulate to such an extent that it causes severe cattle diseases (“locoweed”) (Beyers, 1935) [= Byers, 1935].⁷⁹ In order to explain the action of minute quantities of boron on sugar beets, Brandenburg (1931 or 1939) assumes that a boron atom is placed at the end of a long chain molecule, changing the properties of this molecule.⁸⁰ As to the mode of action of the rare elements we are entirely in the dark. What surmises seem plausible are mentioned in the text.

Lithium. Good cigar contains Li (Bunsen).⁸¹ Young (1935) 110 𝛾/L [10^-6 g/litre] in seawater.⁸² In some instances, seems to be able to replace other alkali metals (Gellhorn).⁸³

Boron. Heart rot of sugar pest. Top disease of tobacco fertility fruit trees. Growth of pollen tubes cocoa. 4.7 mg/litre in seawater (borates change drive of CO₂ in seawater). In mineral boracite Cl₂Mg₆B₁₄O₂₆. In salt 0.89 % B₂O₃. Palaeozoic slates only 0.1 %. In the ash of plants 0.5-1 % B₂O₃. Anions of seawater, according to Errena, not out of eruptive rocks (for Na/Cl, in ocean 1.05/1.09, in erythrocytes 285/4.8; or 0.5 to 59.4). Boron probably from volcanic exhalations.

Fluorine. In Trichoderma koningii (Niethammer).⁸⁴ All iron in seawater as fluoride. Bad effect on teeth. Seawater 1.4 mg/litre. Klement (1938) has shown marine anemones more fluoride in bone and teeth.⁸⁵

Titanium. In nature as ilmenite FeTiO₃ and its weathering product, titanite CaTiSiO₅. Further pseudobrookite, peridotite etc. Rutile is TiO₂, anatase and brookite same composition.

Vanadium. In primitive Chordates (Tunicates). In seawater 0.3 𝛾/L [10^-6 g/litre]. In asphaltic ash (Longobardi). Large amount in Amanitamuscarina (according ter Meulen, see Edelman, 1937).⁸⁶

Clarke (1916, Data of Geochemistry) cites much older literature (p. 712).

Baskervill in ashes peat N. Carolina.

Musingaye in ash coal and oil bearing shales.

Bertin in ashes of plants.

Byle, in lignite from San Rafael, Argentina. Ash 38.22 % V₂O₅.

Momlot, in similar coal, 38.5 % V₂O₅, Torrico and Meck, from Yanli, Peru 38 %. “Grahamite” (oxokenite) from Oklahoma, Wills found 12.2 % V₂O₅ in ash. Nevada asphalt 30 % V₂O₅ in ash.

“Plants have played some part in the concentration of vanadium” (also uranium, see Clarke,1916, p. 717).

Cabriolite = uranium, vanadrite. Vanadium found in fossil plants (Boutwell).⁸⁷

Manganese. In enzymes?

Redox- classical oat disease.⁸⁸

Cocoa, tea and coffee are Mn plants, also rice.

Leo Minder Der Zürichsee im Lichte der Seetypenlehre. Neujahrsblatt Naturforschende Gesellschaft, Zürich, 145 (1943).⁸⁹ Mn accumulation 120 m observations, star shaped colonies ±10𝜇, Leptothrix echinata Beger. No iron at all. “geme𝛽ter Sauerstoffgehalt verbunden mit grösser Vegetationsdichte.” When brought at the air colonies become visible by manganic ion formation. Enormous mass, apparently heterotrophic. When no bacteria present 130 m deep 0.4-0.95 mg/litre. Mn, no iron in vegetation.⁹⁰

Cobalt. In seawater, in rivers, in soils, in salt. Askew has called attention to a curious anaemia in sheep (1936) in New Zealand, which was later observed by Marston in South Australia and also in South Africa. A few milligrams of Co(NO₃)₂ clearly sufficed to cure the disease. It was found that, from certain soils, due to deficiency in Co (or high alkalinity!) the grass contained almost no cobalt.⁹¹ Tried for pernicious anaemia in man without success. Always accompanying iron.

Saporcite CoS, bieberite CoSO₄.7H₂O.

Linnaeite Co₃S₄. However, nearly all igneous.

Nickel. In ash of certain asphalts (Longobardi, 1935). 0.1 𝛾/L [10^-6 g/litre] in seawater. Accumulated by unknown flora and fauna (Edelman). Always accompanies iron.

Millerite NiS, polydimite Ni₄S₅

Beyridite Ni₃S₄, morenosite NiSO₄.7H₂O!

However, nearly all igneous.

Copper. In polyphenoloxide 6.29 % Kurbowitz. In haemocyanine 0.17 %. As anti-araemia in Goitre

“lich disease” of cattle. Colonising disease. Oysters 3 g/kg, in seawater 5 𝛾/L [10^-6 g/litre].

Chief ore: native copper, several sulphides, two oxides, and two carbonates. Sulphate also exists.

Pyrolunite is the chief sulphide. There is an indication that it may be found microbiologically from the sulphate, as in zinc. Chemobiological investigation of mine water might yield interesting results.

Zinc. In enzymes. Widely diffused in rocks, ZnS chief in seawater 5 𝛾/L [10^-6 g/litre], ore sphalerite and wurtzite. Clarke (1916, p. 677). In N. St. Louis, Wheeler found massive zinc embedded in lignite where it had evidently been formed by the reducing action of organic matter upon other zinc compounds. In Galena, Kansas, zinc as ZnS precipitated by organic action? Also, goslanite ZnSO₄.7H₂O occurs. (Thiooxidans?)

Fruit trees, citrus fruit.

Arsenic. Raulin (1869) showed that Aspergillus was greatly stimulated by small amounts of As₂O₃ with nutrient medium 15 𝛾/L [10^-6 g/litre] in oceanwater (see Clarke, 1916, p. 695 for natural occurrence in lithosphere).⁹² It may well be that arsenic, like nitrogen may be reduced to AsH₃ by several fungi. In the older literature there are cases described of green wallpaper in which moulds, introduced by and living in the paper hanger’s paste liberated AsH₃, thus causing pathological effects upon the dwellers in the room. According to others, however, certain other organic compounds of arsenic are the cause of this phenomenon. In any case As³⁺ and not As^5-.

Bromine. Alga, up to 1 % dry substance, Antozoa up to 4 %. In the latter case as 3,5 dibromopyridine. From the mantle of the purple snail 6,6, dibromoindigo. The origin of slugs? Seawater 6.0 mg/litre. Bromine is commercially made out of seawater. It is one of the most abundant minimum elements, and might be mentioned as an élément concentrateur only for a few organisms as the bromine extent of freshwater organisms, land plants and higher animals is very small.

Strontium. In radiolarian shells SrSO₄, suborder Actipylea, genera Actinellus, Acanthociasma, Acanthometron, Acanthonia, Amphilonche, Sphaerocapsa, Diploconus (Kudo, p. 138 and p. 871).⁹³

13 mg/L in seawater. Celestine = SrSO₄. A. Koch found them in bitumen ores limetree (Clarke, 1916, p. 581) together with barite several other cases. In ashes of seaweeds, and Vogel found it in corals and molluscan shells. The find in the Radiolaria is due to O. Bütschli (1907).⁹⁴

Molybdenum. H. ter Meulen (1931). In brain, liver etc. Bortels (1930). Azotobacter, later also for Radiobacter.⁹⁵ In enzymes? In seawater 0.5 𝛾/L [10^-6 g/litre]. Perhaps necessary for any N fixation? Principle ore is the sulphide. Molybdenite MoS₂, often associated, in sedimentary limestone. With pyroxene. Calcite, mica, pyrite etc.

[crossed out:] Antimony

Calcium iodate, lanthanite, in Chilean nitre beds (the calcium ion and the iodine ion have similar ionic radiii and may therefore occur in the same lattice, according to the principle of V.M. Goldschmidt).

Iodine. Di-iodotyroxine in thyroid. Seawater 50 𝛾/litre. Sponges, algae, Lamnaria 0.6 % iodide and iodate. Rain water 0.2-5 𝛾/L [10^-6 g/litre], also river water. 70 % from Chilean saltpetre (iodate, got oxidised in NaCl), 30 % from algae. ⁹⁶ Goitre, iodised salt, mountain districts, N. S. Holland, Java. ⁹⁷

Diagnosis (Mulder) by means of fungi, Cu, Mn. Mulder thesis Wageningen (1938)?⁹⁸ Certain fungi ([Aspergillus niger]) need manganese and copper for their development. By careful experimentation the author has elaborated a method by which pure strains of the fungi are used as indicators in quantitative determination of traces of both of these metals. The method could be extended to a molybdenum determination by means of Azotobacter! etc.

Vernadsky (La Géochimie) speaks about “explosions vitales”, vital explosions.¹⁰¹ When population growth remains a few generations longer in the logarithmic phase (see Sections 4.1.3.and 1.4) these explosive plagues may result. Why is the increasing, or stationary natality not checked by increasing mortality? There may have been abundance of a factor which is a usual minimum factor, there may be an absence of influence inimical, as parasites or consumers, there may have been a combination of the two. But one thing is certain, periodically the locusts come, and the rust, and the plagues and the red death, and perhaps, like the lemming, we, ourselves are recurrent pests, in this day again in explosive ascendency.

Nicolle (1930) [see also Section 5, note 114] has compared the epidemic to an organism. These are necessary for its occurrence a syndrome of phenomena. It starts slowly, slowly it gathers momentum, it breaks all bounds, then it slackens down, remains stationary and, after a few spasmodic flakes often it becomes non-virulent again and dormant in a few carriers.¹⁰²

Epidemies occur in reservoirs with almost maddening regularity. Algae of the Chlamydomonas-type are the most difficult to combat (Whipple). Recently Van Heusden has given a milieu analysis of the water blooms in the Amsterdam Municipal Water Supply.¹⁰³ A classic remains the early work of Hugo de Vries on the organisms that live in the subterranean reservoirs of the Rotterdam water supply (1887). De Vries mentions the occurrence and the role of Bryozoa, Spongilla, and chief of all Crenothix polyspora, an iron bacterium.¹⁰⁴ The epidemies here, as in other cases, are only disturbed links in a cycle – a cycle which, for some reason or other, is temporarily out of its equilibrium. Epidemiology is never the science of one organism by and in itself.

The author had described an epidemic of the centric diatom Aulacodiscus kittoni Arnott at Copalis beach, Washington in 1925. The same diatom was found in masses on the beach in Corinto, Nicaragua, by the author in 1928. Van Heurck mentions the mass occurrence of this species at the mouth of the Congo River. It may well be that it is the abundance of silica after the spring freshlets near the mouth of a large river, which temporarily lifted one limiting factor to the realm of the unlimited. Given sufficient phosphate and nitrate, combined with the enriching influence from the freshwater, the milieu conditions for Aulacodiscus were fulfilled. (See Baas Becking, Tolman, Hashimoto, 1925).¹⁰⁵ Other diatoms, like Melmina, may cause analogous phenomenona (Mare sporco in the Mediterranean).¹⁰⁶

Passed a patrol of luminous jellyfish, spring 1927, near Bay of Tehnantepec in Mexico, 120 miles wide (at least).¹⁰⁷

C. Wiman Palaeont … 1 (1914),¹⁰⁸ 150. Dr W. Sillig, Natur u Museum 57 (1927, p. 94), Russia every 6 years. Pachychilus tartarea and P. migratorius.¹⁰⁹ See also Brehms Tierleben II, curious coin commemorating Anno, 1693 (Fremde Herstrecken in Deutschland gesehen), above: “Ein Diener der Herren der Herrscharen.”

Samuel 6: 5. Philistines vs. David, Golden mice when they sent the ark back.¹¹⁰

California, Wieringen, followed by enormous quantities of birds of prey.¹¹¹ Mice may follow grain, or bumblebees or both.

[Former island] Urk in 1934. The spider plague after the closing of the Zuyderzee may have followed the Chironomus epidemic which occurred because there was no bottom fish to clear the larvae (blood worms). There were no bottom fish because the water was fresh and the bottom salt, so no aquatics available for the oviposition.

Swarm of butterflies the author saw near Palo Alto, California in 1925. The swarm occurred in June, was about 10 miles long and consisted of Vanessa’s, Papilio, Pieris.¹¹² In Java the author witnessed mass occurrence of Libellolids (Aeshna – like things) on the slopes of the Merapi (1936). For two hours (±30 miles) the swarm was observed.

See Baas Becking (1931c), Salt and Salt Manufacture.¹¹³ There are many causes for red water. We name Oscillatoriarubescens, Haematococcus pluvialis, Dunaliella salina, red bacteria (Micrococcus morrhuae Klebahn, 1919), purple bacteria. It is hard to tell whether a freshwater form is meant in the Egyptian plague.

Shelford mentions fly plagues. In the Australian desert [in 1936] the author collected on the head of his assistant [= Dr. J. Reuter] more than 1600 flies.

Especially in the arctic summer, are a well known recurrent nasty epidemic.

(Charles Nicolle Naissance, Vie et Mort des Maladies Infectueuses).¹¹⁴

Lotka.¹¹⁵

5.10.13.a Typhoid

[Baas Becking left this section blank.]

5.10.13.b Malaria

[Baas Becking left this section blank.]

5.10.13.c Cholera

[Baas Becking left this section blank.]

5.10.13.d Influenza

[Baas Becking left this section blank.]

5.10.13.e Plant diseases

[Baas Becking left this section blank.]

5.10.13.f Specific malice

[Baas Becking left this section blank.]

5.10.13.g Mass death (“explosion mortelle”)¹¹⁶

The four horsemen of the Apocalypse (Rev. 6) famine, war, epidemics, deaths.

Suicide (lemmings).

Swarming of termites, drought, euxenic phenomena.

F. Trusheim, Massentot v. Insekten. Natur u Museum, 59, 55 (1929).

Chiefly Lochmaea suturalis Thomas, yellow leaf beetle 2-V-29 Wilhelmshafen [tidal] worms towards dunes then out, cooler, man death 3000 litre, 40 × 10⁶ beetles.

C. Wiman, Palaeont. Zeits., 1, 150 (1914), 150. Dutch steamer in 1899, 33. Locusts Red Sea through counts (2-300/m². D… Pieris, Lüden. Melvl… tha. V. Freyberg Naturw., 15, 13 (1926), Mar Chiquita [Argentina] grasshoppers as salt sea ½ × 15 cm ridge (like Artemia eggs). Dr. Gy Eberle. Ein Massensterben v[on] Heringen, … plant poisonous gas, March 15-16, 1927, Lübeck-Travemunde. Natur u Museum 59, 64 (1929).¹¹⁷

Summarising we might schematise the influences milieu factors by selecting a number of them (Table 5.3a) and classifying the organisms accordingly (Table 5.3b). The relation to the organisms is given as follows:

Of course, the requirements could be further refined, until finally a recipe book for the mass culture of organisms should result. However, this is outside the scope of this essay.

Hippocrates in his classic From the Water and the Places, calls attention to this substance as milieu factor for our living beings: Man. To describe the role of water as a milieu factor would be the same as to write a textbook of physiology, as water enters into metabolism. In this section, however, we only point out a few instances in which water plays a role, such as situations in which water may become “limiting factor.” (Drought, frost, physiological drought, tidal exposure, atmospheric plants) as well as the influence of water upon the shape of organisms, the morphogenetic role of water. Atmospheric moisture, particularly in connection with temperature, has a profound influence on animals, particularly mammals, as shown in Section 5.4.5 for man and sheep.

Have, in their effects, much in common. In both cases water is removed from the cell and the cell solution becomes much more concentrated. “Gefreien and Erfrieren” [Freeing and freezing] (Molisch).

“Wilting coefficient.”¹¹⁸

Dr. J. Zonneveld, from our laboratory (1934).¹¹⁹

Fucus platycarpus.

Fucus serratus.

Fucus vesiculosus.

Ascophyllum nodosum.

Walter (Der Hydratur der Pflanze),¹²⁰ has given a table in which the vapour pressure is given together with the osmotic pressure corresponding. A saturated salt solution lowers the vapour pressure to 85 %, this should be about the limit (tables). MacDougal (1924) and Peirce (1901) have called attention to the redwood Sequoia sempervirens which in California, occurs in the so called “fog chamber”, valleys in which the fog is drawn inland.¹²¹ As a matter of fact, sequoia absorbs water vapour by the leaves, and, maybe there are a great many other plants that do so, only literature is controversy.

Atriplex vesicaria, a chenopodiaceous plant from S. Australia (J. Wood), possesses a perfect root system which it only can use a few weeks in the raining season. The rest of the year it takes up water as vapour by the leaves and Wood obtained values here far above those of Walter and corresponding to swelling pressure of ±1000 atmosphere as the leaves were able to absorb water when the saturation was only 65 %!¹²²

Trentepohlia (Renner). Especially in Java there are a great many species of the beautiful epiphytic or epilithic alga. On the whole, they are unable to take up water vapour when the atmosphere shows less than 85 % saturation. It is a mooted question whether contact with liquid with similar vapour tension creates comparable conditions. It is remarkable that a salt alga Lochmiopsis seems closely allied to Trentepohlia and that also the polyblepharid alga Dunaliella may be a fixed unicellular stage of one (or either) of them (Walter).¹²³ Also, dunalisation [?] occurs in both forms (see Section 7.6.7).

Certain Lichens are able to form where hardly ever liquid water, either dew, or rain, may be found. They apparently are able to absorb water vapour. Goebel has called attention to the enlarged hyphae of the fungeous component (Quellhyphae) which should act as a water absorbing tissue. Quispel however, experimenting with lichens proved that the fungus component did not act as a water reservoir or a water absorbing component.¹²⁴

Higher aquatics …

The redox potential, against a normal H₂ electrode is:¹²⁵

(Table of dyes and substances, work of Elena on redox of surroundings).

Oxidation is:

• 1) addition of oxygen 2H₂ + O₂ = 2H₂O

• 2) removal of electrons Fe²⁺- (e) = Fe³⁺

• 3) removal of hydrogen H₂X + Y = H₂Y + X

Two pages are too short even to mention the multitude of questions which arise here. In Section 5.12.4, where the oxygen balance is dealt with, the problem will be treated more fully.

Pasteur has aptly said “la fermentation, c’est la vie sans air.” Anaerobiosis gives air to fermentation. The products of catabolic aerobiosis are only CO₂ and H₂O, while, in fermentation a whole spectrum of substances may be found. Therefore, in a meristem or at the places where differentiation occurs, the oxygen tension should not be too high (Ruhland) lest the costly metabolites oxidise too far. Maybe that buds in which differentiation takes place often a year before budding (Suringar), are for this reason often protected by wax, resin, and involucrated bracts.¹²⁶ It is remarkable that the analogue of the two chief animal oxygen carriers, the iron containing haemoglobin and the copper-containing haemocyanin, to wit the Fe-containing peroxydes and Cu-containing polyphenoloxydes, occur copiously in plants, in which play other with regulation and mediation like the diënoles, they regulate the oxygen transfer to the substrate. We should not forget, however, that, apparently, biochemistry is not much concerned with oxygen. Hydrogen being the chief component in organic compounds. The oxygen comes in as a sort of afterthought. It may be (see Section 6.4.2.a) that oxygen is really an evolutionary afterthought and that many organisms that once throve upon this earth (Equisetes, Lycopods), all organisms with open and unprotected vegetation points are really suffering nowadays from a sort of surfeit of oxygen. For many anaerobic bacteria oxygen is an active poison, which cannot be withstood in the vegetative state. Transfers of anaerobes should be made, therefore, under special precautions.

Senkenbergiana 14, 1932, Franz Hecht, Der Chemische Einfluss Organischer Zersetzungsstoffe auf das Benthos [dargestelt an Untersuchungen mit marinen Polychaeten, insbesonderes Agricola marina L.], p. 199-220 Arenicola may live in high H₂S without oxygen.¹²⁷Nereis even better. Dissociation curve, Barcroft and Barcroft, The Blood Pigment of Arenicola. Proc. Roy Soc. 96, 192 (1924).¹²⁸

[Baas Becking inserted Fig. 5.25, from Barcroft and Barcroft (1924, p. 36).]

“The difference seems to accord with the differences of function of the haemoglobin in the two forms of life”.¹²⁹ In the higher mammals its duty is to discharge a large quantity of oxygen whilst traversing a capillary in a few seconds of time, in Arenicola the object of the haemoglobin is evidently to return or store oxygen on which the organism can draw back when it is sealed up in its hole at low water.

Milieu study Hecht proves that decomposition benthos may create anaerobic conditions as well. At low water Arenicola usually plenty oxygen. Aerobic lives on its glycogen. The principle of Barcroft might be extended to other respiratory pigments, such as haemocyanin. Plants may grow with their roots in the black mud, stone, like the willow (Cannon) probably transport oxygen chemically. Others transport it directly, as in rice (van Raalte) when it is even excreted.¹³⁰

A disturbed cycle, where, apart from an almost complete exhaustion of oxygen, the H₂S tension is high (Black Sea as example, Walfish Bay).¹³¹ H₂S is toxic as such, in highly alkaline solutions, where chiefly SH^- and S^2- occurs, it is much less harmful. This is perhaps the reason why purple bacteria, that live on H₂S and its dissociation products, only occur at high pH. The significance of an ocean “sill”, see Hardon, Kuenen, Snellius expedition.¹³²

[Explanation of numbering in Table 5.4.]

1) Plankton = 5 ton/Ha, vegetation to 30 ton Ha. Average over earth surface ±10 tons/Ha of which ±30 % carbohydrate, (per annum). 2 ton carbohydrate has liberated out of CO₂ 2.4 ton of oxygen/Ha. 1 Ha = 10⁸ cm², 1 ton = 10⁹ mg, 24 mg/cm² oxygen.

4) According to Clarke (1916) 300 × 10⁶ tons of sulphate disappear into the ocean. They should be reduced. This would make available, for other oxidation 200 × 10⁶ tons of oxygen.

5) Similar order of magnitude.

8) A fraction of 4). Total R₂O₂ to be reduced only 75 × 10⁶ tons.

9) In combustion of coal and oil (±3800 × 10⁶ ton). 4000 × 10⁶ ton of oxygen are absorbed corresponding to 2 mg/km² of the earth’s surface.

Now everything is uncertain, except the order of magnitude per m² there is 7.5 kg C (Neale from Clarke, 1916) in atmosphere per cm² 115 g C and 300 g O₂. Despite the analyses of Goldschmidt, we are very far yet from the construction of a reliable oxygen balance (see Fig. 5.26 and Table 5.5). The humus (50 % C, over 1/3 earth surface 1 %) would amount to 330 mg/cm² carbon (for a layer 1 metre deep). If annual accretion would amount to 1 % of this amount 3 mg/cm² carbon would be shielded from oxidation or a sum of 8 mg/cm². Man consumes on the average ≏ 2000 cal = 3 × 676 cal = 3 × 6 × 32 g O₂ = 576 g O₂. There are 2 × 10⁹ of the species 1.15 × 10¹⁵ mg N per cm² 5 × 10¹⁴ mg. Oxygen.

Now of the total 300 g O₂/cm² (not 230 g, as Goldschmidt calculates from volume % instead of weight %) 10 % enters into the cycle of matter, less than 1 % is due to human activity. Goldschmidt ascribes a very important role to the oxidation of inorganic compounds (weathering of Fe²⁺, of sulphides, of sulphur etc.). Except respiration the only other factor of importance that enter in are 1) the carbon in the humus, and 2) this oxidation of inorganic compounds, of course. The credit side has to be little higher than the debit, the gain, figuring on 1000 × 10⁶ years to accumulate 3 × 10⁵ mg O₂/cm², would 0.0003 mg O₂/cm² of the same order of magnitude as the total respiration of mankind.

The milieu represents a finely balanced set of conditions which, when upset may cause a profound influence upon the vitality, whether in a positive or in a negative sense, of the organism. Most organisms are entirely and passively a prey of the milieu, only in some cases there are regulatory functions (heat regulation; see Section 4.2.5.b). In many instances there is a very narrow relation between organisms and environment, there is a marked resonance of the living being and “le monde ambiant.” Changes in the life cycle, as we shall see, are often caused by changes in the outer world, whether diurnal or seasonal, or both. These influences are super imposed upon those of the internal milieu.