### 5.1.1 Introduction, the “overhanging“fate”

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

### 5.1.2 Nature of the growth curve2

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

$y=ex−e−xex+e−x$

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

$y=11+e−x$

and we write:

$z2=11+e−x−12$

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

$y=ex−e−xex+e−x$

For the Robertson curve:

$y=11+e−x$

we determine the first derivative as

$dydx=e−x1+e−x2=11+e−x⋅e−x1+e−x=y1−y$

the second derivative gives:

$d2ydx2=e−x−e−2x1+e−x3$

the third derivative:

$d3ydx3=e−2x−4e−x+11+e−x6$

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

$e−2x−4e−x+1=0$

or,

$e−x=2±3$

$x=−log2±3log e$

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

### 5.1.3 Consideration of H. Mark on the growth of large molecules

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!

### 5.1.4 Derivation from normal, ogive and other expressions

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

$y=1σ2π⋅e−x22σ2$

is an example, the integral of:

$y=axax2+bx+c$

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

### 5.1.5 Examples of natural growth curve, chiefly of populations

[Baas Becking left this section blank.]

### 5.1.6 Analysis

[Baas Becking left this section blank.]

### 5.1.7 Summary and conclusions

[Baas Becking left this section blank.]

### 5.2.1 Introduction

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.

### 5.2.2 The factors rehearsed: properties of water, radiation

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

### 5.2.3 Stenobionts

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).]

### 5.2.4 Eurybionts

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.

### 5.2.5 Summary and conclusions

[Baas Becking left this section blank.]

### 5.3.1 Introduction

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).

### 5.3.2 Photosynthesis

[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.]

### 5.3.4 Vision

Eye of Gyrinus.19 Melanophore contraction.

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

### 5.3.5 Formative influences

Light and shadow lenses. Etiolation.20 Other influences.

### 5.3.6 Photoperiodicity

[Baas Becking left this section blank.]

### 5.3.7 Germicide action21

[Baas Becking left this section blank.]

### 5.3.8 The ultraviolet

[Baas Becking left this section blank.]

### 5.3.9 Germitisation

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

### 5.3.10 Summary and conclusions

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

Text box 5.1 – Baas Becking notes made prior to writing the manuscript

Baerends (1943) shows how little change in the ocean water sufficient change composition fish fauna, tunny not <14 °C, etc., sole etc.22 Schenk (1917), Interlaken,23 temperature of 200-300 °C in heating hay! (see Schwarz and Laupper, 1922).

### 5.4.1 Introduction

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.

### 5.4.2 Influence of very low temperatures

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

### 5.4.3 Influence of very high temperatures

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

### 5.4.4 Temperature table

[Baas Becking left this section blank.]

### 5.4.5 Temperature and humidity

[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).

### 5.4.6 Summary and conclusions

[Baas Becking left this section blank.]

Text box 5.2 – Baas Becking notes made prior to writing the manuscript

Protozoa (Kudo) including authors. Euglena spp. pH 3.0-9.9, Paramoecium sp. pH 7.0-8.5, Paramoecium caudatum Ph 5.3-8.2, Stylonychia pustulata pH 6.0-8.0, Colpidium sp. pH 6.0-8.5, Colpoda cucullus pH 5.5-9.5.

### 5.5.1 Introduction

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.’

### 5.5.2 pH and pH range of the milieu

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

### 5.5.3 Oxalic acid

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

$COOH2=2CO+H2O2,$

and with access of water

$2CO+2H2O=2 HCOOH$

$2 HCOOH=4 H+2CO2,$

and with access of oxygen

$4H+O2=2 H2O.$

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).

### 5.5.4 Occurrence of mineral milieu as solute as function of milieu

[Baas Becking left this section blank.]

### 5.5.5 Summary and conclusions

[Baas Becking left this section blank.]

### 5.6.1 Introduction

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

### 5.6.2 Osmotic pressure and osmotic phenomena

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.

### 5.6.3 Simulacra vitae

(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 SO42- 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.

### 5.6.4 The rhythm of the pulsatory vacuole

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

### 5.6.5 The secretion and intake of water

[Baas Becking left this section blank.]

### 5.6.6 The problem of salt organisms

[Baas Becking left this section blank.]

### 5.6.7 Theory of salt organisms

[Baas Becking left this section blank.]

### 5.6.8 Theory of water

[Baas Becking left this section blank.]

### 5.6.9 Osmophilic organisms

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

M. Fischer.44

### 5.6.12 Summary and conclusions

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

### 5.7.1 Introduction

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.

### 5.7.2 Historical: Ringer

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.

### 5.7.3 Historical: Loeb, Osterhout46

[Baas Becking left this section blank.]

### 5.7.4 Hypothesis of seawater: McClendon47

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.

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

### 5.7.5 Classification of phenomena

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

### 5.7.6 Natural waters again

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)

• 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.

### 5.7.7 Artemia

[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

### 5.7.8 Oöspora and other microbes

[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[?].

### 5.7.9 Lochmiopsis sibirica Woron ( = Ctenocladus circinatus Borzi)

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.]

### 5.7.10 Dunaliella viridis Teod

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.]

### 5.7.11 Other salt organisms, algae

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.]

### 5.7.12 Pollen grains

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

### 5.7.14 Summary and conclusions

[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).]

### 5.8.1 Introduction

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

### 5.8.2 Teleology

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

### 5.8.3 Terminology

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:

$y=y0σ2π⋅e−x22σ2$

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

### 5.8.5 Higher organisms

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?

### 5.8.6 “Fremddienliche Zweckmässigkeit” [Expediency for other purposes]

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.

### 5.8.7 Mimicry

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).

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 As5-.

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.

Text box 5.3 – Baas Becking notes made prior to writing the manuscript

Beijerinck, later H.G. Derx, Bact violaceum. Slime of bread, next to trickle of the tap!99 Sarcina (see also J. Smit).100

### 5.10.1 Introduction

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.

### 5.10.2 Plagues, epidemies

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

### 5.10.3 Water supplies

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.

### 5.10.4 Diatoms

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

### 5.10.5 Jellyfish

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

### 5.10.6 Locusts

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.”

### 5.10.7 Mice

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.

### 5.10.8 Spiders

[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.

### 5.10.9 Other arthropods

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.

### 5.10.10 Red water

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.

### 5.10.11 Flies

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.

### 5.10.12 Mosquitos

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

### 5.10.13 Epidemiology

(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

### 5.10.14 Milieu chart

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.

### 5.11.1 Introduction

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.

### 5.11.2 Drought and frost

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

### 5.11.3 Tidal exposure

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

Fucus platycarpus.

Fucus serratus.

Fucus vesiculosus.

Ascophyllum nodosum.

### 5.11.4 Intake of water vapour

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

### 5.11.5 Morphogenetic influence of water

Higher aquatics …

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

$E=RTN ln oxred−F$

(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.

### 5.12.1 Anaerobiosis, aerobiosis

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.

### 5.12.2 Senckenbergiana

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

### 5.12.3 Euxinic phenomena (“Verjauchzung”) [= Putrefaction]

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 S2- 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

### 5.12.4 Processes which influence the terrestrial oxygen balance

[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.

1
Baas Becking probably referred to Pearl (1926, The Biology of Population Growth). Raymond Pearl (1879-1940), American biologist and T.B. Robertson, a physiologist. He published in 1908 two articles applying the logistic curve to various cases of individual growth in animals, plants and man (Robertson, 1908). The logistic curve was introduced by Raymond Pearl and statistician Lowell Reed in 1920, unaware of Robertson’s earlier work. Pearl promoted the sigmoidal curve as a description of human and animal population growth. In subsequent years it underwent a barrage of criticism from statisticians, economists, and biologists, a barrage directed mostly against Pearl’s claim that the logistic curve was a law of growth. Nevertheless, it emerged in the mid-1930s as a central model of experimental population biology, and in its various modifications has remained an important part of modern population ecology.
In the 1953 version of Geobiology, Baas Becking (1953a) discussed the work of Raymond Pearl in Chapter VIII, Section 10 Human Population, p. 754-759.
2
Baas Becking (1946a) described a number of sigmoid curves and their application to data of growth of pumpkins, rats and yeasts, growth of baby’s (weight) and man (length) and population increase. The cases are described, the biological comments are not summarised in the final conclusions. He wrote about sigmoidal growth:
Unless organisms have second sight, it is hard to conceive how the sigmoid should possess a predetermined upper limit for y or, in other words, when we do not want to accept an entirely predestined milieu, the sigmoid, as originated from the logistic expression, loses its biological meaning.
He also stated that
Goodness of fit of a certain expression to a certain set of experimental data does not mean that those data actually are connected by a law which is expressed by the equation of the curve which is tried.
In the 1953 manuscript of Geobiology Baas Becking wrote a comprehensive Section on Organisms as a Function of Time (p. 118-133).
3
A reference to Herman Franz Mark (1895-1992), Austrian chemist, ‘father of polymer science’, who escaped from Nazi Europe in 1938 and came to the Polytechnic Institute of Brooklyn. Baas Becking referred to The General Chemistry of High Polymeric Substances (Mark, 1940). The book was discussed by G. Salomon in Chemisch Weekblad, (February 8, 1941, p. 68-69). July 5, 1944, Mrs. T. Niekerk-Blom supplied him with a copy in the Utrecht Kriegswehrmachtgefängnis (NIOD 214, nr 33).
See Morawetz (1995), Deichmann (2001, p. 181-187).
4
Baas-Becking and Baker (1926).
5
Corpora non agunt nisi fluida (or liquida) seu solute (Aristotle): Compounds do not react unless fluid or if dissolved.
6
Like in Section 3.5.1, Baas Becking referred to Rev. William Whewell (1794-1866).
7
In his lecture for the Sydney University Biological Society The Nature of Death, July 14th, 1953, Baas Becking also quoted Robert Seymour Bridges (1844-1930). He referred to The Testament of Beauty: a Poem in Four Books (Bridges, 1929).
Life is an improbable event. As Robert Bridges says; “it holds balance on a razor’s edge, that may not e’en be blunted, lest we sicken and die.”
8
Reference to Johannes Carolus van der Meer Mohr (1892-1969), entomologist, in 1925 assistant, and from 1934-1939 director, Deli Experimental Station Medan. Van der Meer Mohr (1929), Van der Meer Mohr (1931).
9
Reference to dissertation Jacoba Ruinen (1933) in which the species is named Lochmiopsis siberica Woron. This form is identical to Ctenocladus circinnatus Borzi. See also Baas Becking (1934 and 2016, p. 117).
10
Vernalisation is the induction of a plant’s flowering process by exposure to the prolonged cold of winter, or by an artificial equivalent. In 1928 the Russian agronomist Trofim Lysenko coined the term ‘jarovisation’ to describe a chilling process he used to make seeds of winter cereals behave like spring cereals. Later Lysenko inaccurately asserted that the vernalisated state could be inherited. During the 6th International Botanical Congress in Amsterdam in 1935 the Russian work on ‘jarovisation’ was discussed (Proceedings v. I, p. 158-160):
There seems to be complete agreement in Russia as to the value of the method and it is there hailed, as one of the greatest advances in crop production yet made. Experimenters in other countries, however, have been much less enthousiastic and there is wide divergence of opinion as to its effectiveness.
Trofim Denisovich Lysenko (1898-1976), Soviet agronomist and biologist, responsible for subsuming biology to ideology. According to Andrei Sakharov in 1964:
He was responsible for the shameful backwardness of Soviet biology and of genetics in particular, for the dissemination of pseudo-scientific views, for adventurism, for the degradation of learning and for the defamation, firing arrest, even death, of many genuine scientists.
Apparently, Baas Becking was unaware of the scientific controversy over Lysenko’s ‘vernalisation’experiments. In his 1943 notes for Geobiology (1944) he summarised Lysenko’s 1928 experiments (AAS 043 nr. 160-6):
VERNALISATION applies to temperate wheat plants. Vernalisation is a technique by which flower production and fruiting is accelerated by application to the germinating grain if temperatures slightly above freezing point. […] Lysenko introduced his idea of “phasic development” of plants. […] The phenomenon of vernalisation is more complex than this and masks the focal point of two quite separate lines of investigations. One derived from Gassner’s work on influences of temperature during germination; the other from the photoperiodic studies of Gardner, Allard and of Rasmund. Our knowledge of vernalisation has been advanced chiefly by the studies of O.N. Purvis.
O.N. Purvis and co-workers published six studies on vernalisation of cereals in the Annals of Botany 1934-1939. In his 1953 manuscript of Geobiology Baas Becking still quoted Lysenko as an authority (p. 430-431):
The systematic investigation of periodic temperature sensitivity has led to the so called vernalisation of winter grains. These grains are normally sown in the autumn, as they need a cold period for the formation of flowers in the spring. Summer grains are sown in the spring and form ears without a previous cold period. Purvis (1939) has shown that summer rye differs in one dominant gene from winter rye. It is well known from work of Lysenko that winter wheat may acquire the characteristics of summer wheat if the seeds are subjected to a temperature of 3-10 °C for a few weeks. This vernalisation maybe applied to many other plants e.g., potatoes, grapes, soybeans, vetches, maize, rye, red clover and sunflower. (Vernalisation Bulletin 1935).
See Purvis (1961) for review of his vernalisation studies, Gassner (1910), Gardner and Allard (1920).
11
Reference to PhD dissertation Jacoba Ruinen (1933; Figure 17, p. 780-781):
The temperature range in which akinetes show rapid and intense germination is very narrow. The lowest limit lies at 14 ° C. At this temperature germination is reduced to a few percent, while growth has almost stopped. An optimum was found at 18 °C. Here germination occurred after a few days, the young plants were almost 0.5 mm. long. Growth seems still vigorous between 18°-21 °C., but here germination decreases markedly.
12
In Geobiologie (1934) Baas Becking referred to this finding in his laboratory in Leiden (p. 50, Eng. edition, 2016).
13
Reference to Gordon Floyd Ferris (1893-1958), American entomologist, Professor of Biology, Entomology at Stanford University (1912-1958). Baas Becking referred to Ferris (1916).
14
In the 1953 version of Geobiology Baas Becking wrote (p. 398-399):
The most acid volcanic water on record is that of the Kali Pait, on the slope of the Idjen volcano, Java. Here no less than 216 mg H+ was found per litre, corresponding to a pH of 0.66. the acid was […] mainly HCl. It should be remarked that certain bacterial sulphide oxidation may cause even higher acidities.
15
In the 1953 version of Geobiology (p. 399), Baas Becking referred to the tolerance of Amoeba in the hot spring in the volcanic water of the Javan Patuha:
The high acidity seems to be no limiting factor either. Kawah Tjiwedéh on the slope of the Patuha in Java (temp. 42-85°C), although the pH was below 2.0, was teeming with life, even an amoeba being present (see Ruinen and Baas Becking, 1938).
The reference to Kawah Tjiwedéh is also in Baas Becking (1938a), On the Cause of the High Acidity in Natural Waters, especially in Brines.
17
In the 1953 manuscript of Geobiology (p. 393-397) Baas Becking described the case of Lake Bumbunga.
18
This is evidently a note from Baas Becking to remind him that this section needs to be expanded.
19
Reference to the whirligig beetle, Gyrinus substriatus or Gyrinusnatator.Xenogyrinus natans, extinct species, well known in Dutch literature as ‘Het Schrijverke (Gyrinus natans)’ by Guido Gezelle. This species was first described in 1845 by Peter Bellinger. However, Gezelle referred to the before mentioned species G. substriatus or G. natator.
20
Etiolation is a process in flowering plants grown in partial or complete absence of light. It is characterised by long, weak stems; smaller leaves due to longer internodes; and a pale yellow colour (chlorosis). Elongation is controlled by auxines which are produced by the growing tip to maintain apical dominance. Auxin diffuses, and is transported, downwards from the tip, with effects including suppressing growth of lateral buds. Auxins are active in light; when they are active, they stimulate proton pumps in the cell wall which increases the acidity of the cell wall and activates expansine (an enzyme that breaks bonds in the cell wall structure) that weaken the cell wall and allow the cell to expand.
21
Baas Becking probably referred to Ultraviolet Germicidal Irradiation, a means of disinfection. The bacterial action of ultraviolet light was studied in the 20s and 30s of the last century.
See Reed (1974), The History of Ultraviolet Germicidal Irradiation for Air Disinfection.
22
‘Baerends (1943)’ not identified. Reference to Gerard Pieter Baerends (1916-1999) Dutch ethologist and fisheries biologist, Professor of Zoology at Groningen University.
See for his relationship as a student in Leiden with Baas Becking: Baerends (1985).
23
Not identified.
24
Baas Becking referred to Bastiaan Jacob Dirk Meeuse (1916-1998). During his study in Leiden, Meeuse was inspired by Baas Becking. The reference is to his 1943 PhD thesis, Oriënterende Onderzoekingen over de Vorming van Rietsuiker uit Zetmeel in Planten bij Lage Temperatuur (Delft, G. van Iterson, supervisor). From 1952 until 1986, Meeuse was Professor of Botany at the University of Washington, Seattle.
26
Reference to the Russian-British entomologist David Keilin (1887-1963), most known for his research and re-discovery of cytochrome in the 1920s: Keilin (1925). See also Slater (2003).
27
‘Proteid’ is an obsolete word for a protein molecule.
28
Reference to Gortner (1930). In the 1953 version of Geobiology Baas Becking referred to Gortner:
The problem of the state of water in colloidal systems was, in the meanwhile, approached from another angle. Gortner approached the problem from a colloid chemical angle, that more advance was made. According to Gortner, water may exist, in a colloid (like protoplasm) in various states: it may exist as free molecules, as molecules bound osmotically or as hydrates and further it may occur as swelling water, tightly bound to hydrophilic, colloidal particles.
29
Reference to Baas Becking (1942b). See also Section 3.5.21, Free and bound water.
The section about deep freezing fruit was probably also based on the work of Baas Becking and Henri Derx for Unilever (1942-1944). See for reference to the classified reports Bibliography L.G.M. Baas Becking and of Pupils and Co-workers 1948. Document AAS Basser Library 043 nr. 161. In the 1953 version of Geobiology Baas Becking referred to his research in the Unilever laboratory (see Section 3, note 72).
30
Apparently Baas Becking took the two diagrams from Shelford (1929). The left hand graphs were taken from the work of Ellsworth Huntington (1876-1947), an American geographer known for his studies on environmental determinism. The right hand diagram was taken from Johnson (1924).
31
Dr. P.S.J. Schure, Baas Becking was her PhD supervisor in Leiden. The reference is to Schure (1935a and 1935b), Table 14, Survival of the Swarm Cells in Drops with Few Spores on Different Surfaces.
32
Kazimierz Bassalik (1879-1960), Polish microbiologist, professor Warsaw University. Reference to Bassalik (1913), his ‘Bacillus extorquens n. sp’ has the present name Methylobacterium extorquens. See also Bassalik-Chabielska (1980).
33
In the 1953 manuscript of Geobiology Baas Becking (p. 657) referred to C.J. Niekerk-Blom’s 1946 publication:
The ‘lonely’ compound at reduction level (1), oxalic acid, remains a biological enigma. It is of very wide occurrence in the plant kingdom, and often most abundant, but its mode of origin is still obscure. It is oxidised both by bacteria and by higher plants, sometimes with the formation of hydrogen peroxide.
Mrs Niekerk referred to Zaleski and Reinhard (1911), who discovered the degradation of oxalate by plant material in wheat. She also referred to Bassalik (1913) and to Franke and Hasse (1937).
See for more recent research: Davoine et al. (2001), who investigated changes in the activity of the extracellular enzyme oxalate oxidase and the concentrations of oxalate and H2O2 during the ageing of leaf sheaths of ryegrass. The accumulation of H2O2 during ageing coincides with the increase of both oxalate level and oxalate oxidase activity. Overall, results suggest that in ryegrass that synthesises both Ca oxalate and oxalate oxidase, the production of H2O2 and Ca2+ during ageing of stubble might be involved in the constitutive defenses against pathogens, thus allowing the phloem mobilisation of nutrient reserves from the leaf sheaths towards elongating leaf bases of ryegrass.
34
Reference to James William McBain (1882-1953). Canadian/American chemist. Professor of Chemistry, University of Stanford (1926-1947). He devoted many years to the elucidation of the nature and structure of soap-rich aqueous systems.
35
Erik Zevenhuizen (2008), in his biography of Hugo de Vries, discussed the work of Hugo de Vries, H.J. Hamburger and H. van ’t Hoff on osmosis in cells of plants and animals in the late 1870s and 1880s (p. 172-180). See his notes 191 and 192 on p. 559-560.
36
For ‘recretion’ and ‘intrability’ see Section 4.2.5.a.
37
See Zevenhuizen (2008,), p. 172-175, 559 (notes 187-190). J.H. van ’t Hoff (1884) used the data from Pfeffer (1877) for his Etudes de dynamique chimique in which he published his results of osmotic studies in solutions.
38
In the lemna ‘Protoplasma’ in the Encyclopaedisch Handboek van het Modern Denken (1931/1942), Baas Becking remarked:
Speaking of “living matter” is termed ”logical nonsense” by modern Russian researchers; And in my opinion rightly so. Driven by the desire to make life itself one day in the test tube and specially to put the monadological and monistic crown on the great unifying work of the last century, many have allowed themselves to be seduced into premature conception. The whole interlude of “the living matter” is superfluous in the development of science and would never have taken place if the word protoplasm had been left alone without linking it to the then development of chemistry. As an afterglow of this school one still finds people who, in the imitations of life phenomena, as one shows them in the test tube (the so called simulacrum vitae) see indications of how one will have to prepare the “living matter” in the not too distant future.
Raphael Dubois (1849-1929), Professor of Physiology at Lyon, produced particles that were supposedly identical to bacterial cells by adding barium or radium chloride to sterilised fish broth. The reference is to Dubois (1919).
Alfonso Luis Herrera (1868-1942), Mexican physiologist, was convinced that life could be created in the laboratory. He proposed an autotrophic theory he called plasmogeny. See Cleaves et al. (2014).
Moritz Traube (1826-1894), German physiologist. In 1864 Traube was the first to produce semipermeable membranes. See Traube (1866).
39
Liesegang rings, a phenomenon seen in many, if not most, chemical systems undergoing a precipitation reaction under certain conditions of concentration and in the absence of convection. Rings are formed when weakly soluble salts are produced from reaction of two soluble substances, one of which is dissolved in a gel medium
40
The reference is to Charles Vincent Taylor (1885-1946), an American biologist. His doctoral dissertation, done under the supervision of the parasitologist Professor Charles A. Kofoid, was entitled Demonstration of the Function of the Neuromotor Apparatus in Euplotes by the Method of Microdissection. Taylor was active at the University of California and the Hopkins Marine Station of Stanford University in the 1920s when Baas Becking was working at Stanford and the Marine Station. In 1933 Taylor was made Herzstein Professor of Biology, the chair that was occupied by Baas Becking until 1931.
Baas Becking referred to Taylor (1923), The Contractile Vacuole inEuplotes. Contractile vacuoles are subcellular organelles which are defined by their behaviour of filling slowly with fluid, and periodically expelling their contents from the cell. It is a membrane bound osmoregulatory organelle of freshwater and soil amoebae and protozoa which segregates excess cytosolic water, acquired osmotically, and expel it to the cell exterior, so that the cytosolic osmolarity is kept constant under a given osmotic condition.
41
Jean Paul Louis Pelseneer (1863-1945), Belgian malacologist, morphologist, ethologist and phylogenist. Baas Becking probably referred to Pelseneer (1905). In this study Pelseneer discussed the osmotic changes in fishes migrating from a saline to freshwater environment.
Yves Delage (1854-1920), French zoologist, since 1901 director Station Biologique de Roscoff. Baas Becking probably referred to Delage (1895).
42
The reference to the polychaete worm Nereis and to the American zoologist and palaeontologist William Patten (1861-1932) and his research on the vision of Arca noae and Nereis. See Patten (1886).
43
Evadne refers to marine crustacea that have the ability to stay alive for a long period of time in distilled water. See Baas Becking in Geobiologie (1934, English edition, p. 50). See also Section 5.2.3.
44
Reference to H.W. Fischer (1910), Gefrieren und Erfrieren. See also H.W. Fischer (1911), Das Wasser im plasma.
45
Perfusion is the passage of fluid through the circulatory system or lymphatic system to an organ or tissue, usually referring to the delivery of blood to a capillary bed in tissue. Baas Becking discussed the Ringer experiments in Geobiologie (1934, p. 100-101 in English edition, 2016).
46
Reference to Jacques Loeb (1859-1924), German born American physiologist and biologist. Since 1910 he remained at the Rockefeller Institute for Medical Research in New York. He spent his summers at the Marine Biological Laboratory in Woods Hole. March 4, 1922 Baas Becking wrote to F.A.F.C. Went that he visited Loeb and Loeb’s former assistant, the physiologist and Harvard Professor W.J.V. Osterhout (1871-1964) on his journey from Holland to Palo Alto. Loeb and Osterhout were editors of the Journal of General Physiology. They published two papers by Baas Becking in 1920 and 1921 [Hampton and Baas Becking (1920) and Baas Becking (1921c)].
Letter L.G.M. Baas Becking to F.A.F.C. Went Stanford University Palo Alto March 4, 1922. Correspondence F.A.F.C. Went Library Boerhaave Museum Leiden.
In his 1951 lecture Forgotten Biology, Baas Becking (1951b) referred to Loeb as:
My respected teacher, Jacques Loeb, saw a future for biology in which the certainty of the results would equal those in chemistry and physics. However, the principle of uncertainty has entered in both chemistry and physics and, as living beings represent statistical populations of a much lesser magnitude than those met with in the molecular the atomic or the electronic state. The individual freedom of the variant always leaves its hallmark upon the results. We are striving towards the impossible if we set our course solely by the lights of our neighbouring sciences. Apart from the two attitudes towards biology mentioned above, there are recurrent raves of vitalism and of mechanistic thinking, which cloud the picture.
In 1927 in his course General Physiology of the Cell at Utrecht University, Baas Becking (1927-1928) referred to Loeb:
Physical chemistry has been claimed by many to be the most powerful tool for the physiologist. Sinclair Lewis in his “Arrowsmith” made Professor Gottlieb (obviously Jacques Loeb) say:
Organic chemistry! Puzzle chemistry! Stink chemistry! Dungstore chemistry! Physical chemistry is power, it is exactness, it is life. But organic chemistry – that is a trade for pot washers.”
47
Baas Becking discussed McClendon in Chapter IX, Geobiologie (1934) and copied a figure from his publication, see English edition Geobiologie (2016, p. 96-97). Jesse Francis McClendon (1880-1976), an American chemist zoologist and physiologist, known for the first pH measurement of human stomach in situ. The reference is to McClendon (1916).
The reference to McClendon in the 1944 manuscript of Geobiology is probably a mistake for Macallum. In his Inaugural Address as extraordinary professor at Utrecht University, October 3, 1927, Baas Becking referred to Archibald Byron Macallum (1858-1934), who published in 1908, Cellular Osmosis and Heredity’.
48
In the 1953 manuscript of Geobiology Baas Becking discussed six salt-water mineral wells at Sapta Tirta near Pablengan, Java (p. 378-379):
From the analyses of ter Haar it appears that the salt waters of this region represent a distinct type, resembling seawater, but with less sulphate. The high iodide and ammonia (see analysis Delok) [a salt water well near the town of Tjopu near the river Gagakan] show a large biogenic element in the origin of these waters. Rather close to this type of water is a brackish lake near Rockanje on the island of Voorne, Holland, known for its lime deposits.
The high calcium carbonate content of the water mentioned and the comparatively low pH (ranging from 5.9 – 7.2, average 6.5) together with its often very high content of H2S and suspended lime may be conceived as seawater in contact with groundwater and calcium carbonate with a large amount of sulphate reduction. [Baas Becking inserted: To account for the disappearance of the anion]. The disappearance of the magnesium is not accounted for, unless a permutate action (base exchange) with soil particles might be the cause (Takir Ghiol [Lake Techirghiol Rumania, known for its sapropel: dark coloured sediments rich in organic matter]).
Baas Becking probably referred to Ir. Carel ter Haar, a Dutch geologist who discovered in 1931 along the Solo River on Java fossils of Homo erectus, nowadays considered the last Homo erectus, which lived until 200,000 years ago.
49
The relation between internal fluids of organisms and seawater was a topic of fascination for Baas Becking. In the 1934 Geobiologie, he discussed the physiological and evolutionary significance of this (p. 100-101; Eng. version). In the 1953 manuscript of Geobiology (p. 487-488) Baas Becking wrote:
Macallum, in 1908, considered the blood serum of higher animals as an ancestral seawater trapped, as it were in the milieu interne. Henderson (1913) adhered to this belief. He states (l.c.p. 187) “not only do the body fluids of the lower forms of marine life correspond with seawater in their composition, but there are at least strong indications that the fluids of the highest animals are really descended from seawater.” This statement has been challenged by many (Baas Becking, 1934; Hutchinson, 1948) but it should receive the attention that it deserves as a bold and inspiring. [….] If Macallum’s hypothesis were true, and organisms arose, in the course of evolution, from their ancestral sea, the first emergent creatures would have left an ocean of a very curious composition, the salts consisting chiefly of NaCl, with a rather high calcium and carbonate content, and much lower in magnesium and, especially in sulphate, then the present ocean. We have to account, moreover, for an ever increasing concentration of potassium. It would run counter to the hypothesis of a constant plutonic addition of chloride, but rather strengthen the theory that, when water became liquid, there was already much halite available on the earth. [Baas Becking inserted: which salt was dissolved first.] Dr. G.W. Leeper called my attention to the fact that NaCl melts at 800 °C and could, therefore, have occupied basins on the earth’s surface when the first rocks solidified. It must have been, (if we take the consequences of Macallum’s theory) a chloro-carbonate to a chloride water of approximately 0.7-0.9 % NaCl, faintly alkaline and in composition not unlike Natron Lake, near Thebes, Egypt (total solids 0.441 %). Much of the charm of this speculation, however, disappears when we take into account the osmo-regulative powers, especially in the higher vertebrates. A similar argument as given above might be applied to the salt content of the blood, in order to arrive at the concentration of an archaic seawater.
Macallum (1926) noted that, although similarities between seawater and organismal fluids, such as blood and lymph, indicate that the first animals emerged in the sea, the inorganic composition of the cell cytosol dramatically differs from that of modern seawater. Macallum insightfully pointed out that “the cell […] has endowments transmitted from a past as remote as the origin of life on earth.”
Baas Becking further referred to Geoffrey Winthrop Leeper (1903-1986), from 1946 till 1962 Associate Professor in Agricultural Chemistry at the University of Melbourne. In 1962 professor. Leeper edited The Evolution of Living Organisms (1962), with papers of the symposium to mark the centenary of Darwin’s Origin of Species, held in Melbourne, December 1959, which contained Baas Becking’s paper On the Origin of Life.
50
Baas Becking referred to Jacques Loeb’s experiments on the role of salt for preservation of life. Loeb (1911), Loeb (1914).
In the 1953 version of Geobiology Baas Becking referred in Section Antagonism on p. 483 to Loeb and Osterhout:
Especially through the work of Loeb and Osterhout a great many cases of this antagonism, both in animals and in plant have come to light. Mixtures of salts which antagonise completely the individual toxic actions are called “balanced solutions.” Ringers solution, widely used in physiological practice, is such a balanced mixture. In general, the antagonistic effect is most pronounced if the valence of the component ions is different. It has been found that NaCl and CaCl2 solutions, both slightly hypotonic for plant cells cause plasmolysis when mixed in a proportion 10 NaCl : 1 CaCl2. It appeared that the solutions, containing the single salts, increased the permeability of the protoplast while, in combination the permeability was decreased, so that plasmolysis could take place. The permeability of the protoplast increases by application of cations in the following sequence:
Ca < Ba < Mg < Li < Na < K
[…]
Loeb has already shown that seawater could be diluted over a rather large range, without a change in physiological effect on various organisms, if the ionic proportions remained the same. The influence of ionic proportions may be studied to great advantage in euryhaline organisms, as here the organisms seem, in a way, independent of its osmotic milieu. […]
Loeb has sung the praise of seawater as a balanced solution. Blood is certainly a balanced solution. It seemed inevitable that these two should be brought together. In 1902 G. Bunge (cited by Lotka, 1924, p. 203) expressed the opinion that be salt present in vertebrates, especially in the cartilage and in the serum, might be interpreted on an evolutionary basis.
51
Reference to Boone and Baas Becking (1931).
52
Reference to Jacobi and Baas Becking (1933).
54
The explanation was taken from paragraph “ionic antagonism” in the unpublished essay Seawater as a Chemical Milieu (Baas Becking, 1945-1946). Oren (2011, p. 21-22) discussed Baas Becking’s experiments on the limits of salt concentrations and other environmental conditions with special emphasis on the antagonistic effects of different ions.
55
See for the 1935 dissertation of T. Hof (Hof, 1935a) also Section 5.2.4.
56
Baas Becking referred to his Dunaliella experiments which he presented at the Hopkins Marine Station in December 1929 at the Midwinter meeting of the Western Society of Naturalists (Baas Becking, 1930). The results of his experiments were further published in The Journal of General Physiology (Baas Becking, 1931a) and in The Scientific Monthly of May 1931 (Baas Becking, 1931c). See Oren (2011, p.13-17) for a discussion of Baas Becking’s experiments with Dunaliella.
57
Reference to Jetske de Zeeuw (1939).
58
Possibly reference to Jacob Reitsma (1932), who defended his PhD dissertation, Studiën über Armillaria Mellea (Vahl) Quél. in Utrecht (supervisor Johanna Westerdijk). Also reference to Booij (1940), The Protoplasmic Membrane Regarded as a Complex. PhD Thesis, Leiden. H.L. Booij defended his Leiden thesis in 1938 (supervisor H.G. Bungenberg de Jong), it was published in 1940.
59
Reference to Theunissen (1936), Lyophiele niet Amphotere Bio-kolloiden Beschouwd als Electrolyten. Thesis, Leiden. Theunissen’s work and that of his teacher Bungenberg de Jong and colleagues H.L. Booij and J.G. Wakkie in Leiden, was concerned with the reversal of charge of biocolloids with salt mixtures. They studied the phenomenon in physiology that some cations may antagonise the action of others. In the 1953 version of Geobiology (1953a, p. 284) Baas Becking referred to:
Dr. J. Wakkie has shown in the Leyden Laboratory (1932, unpublished) that various simple Chlorophyceae are sensitive to cations in low concentrations.
Baas Becking co-authored: Bungenberg de Jong, Vander Meer and Baas Becking (1935), Kolloidmodelle zur Illustration Biologischer Vorgänge I. Dreisalzeffekte bei der Keimung von Krustentiereiern und bei Phosphatiden.
60
Michael F. Guyer (1874-1959) and Elizabeth A. Smith performed experiments attempting to demonstrate Lamarckism (Guyer and Smith, 1920). Their experiments were criticised and were not repeated by other scientists.
61
Paul Kammerer (1880-1926) Austrian biologist advocated Lamarckism. He was accused of fraud for his experiments with midwife toads. Six weeks after the accusation Kammerer committed suicide.
62
In 1942 Krythe and Wellensiek of the Institute for Plant Improvement, in Wageningen the Netherlands, were able to report that polyploidy had been induced by colchicine in 243 species. After reviewing the literature from 1937, they were able also to state, however, that the results of artificial polyploidy had been largely over confident and over optimistic. Krythe and Wellensiek (1942). See also De Chadarevian and Kamminga (2005).
63
‘Ignoramus et ignorabimus’, meaning ‘we do not know and will not know’: scientific knowledge is limited.
64
Gottlieb Haberlandt (1854-1945), Austrian botanist who wrote Physiologische Pflanzenanatomie, published in 1884 (Haberlandt, 1884). This book inspired Hugo de Vries. Haberlandt combined the results of the physiological processes of organs, where physical and chemical laws are guiding, with the interaction with the anatomy and function of organs. So, he made a difference between processes in the animate and inanimate nature. See Zevenhuizen (2008, p. 182-183).
65
Hans Driesch (1867-1941), German biologist and philosopher. Driesch, believing that his results compromised contemporary mechanistic theories of ontogeny, instead proposed that the autonomy of life that he deduced from this persistence of embryological development despite interferences was due to what he called entelechy, a term borrowed from Aristotle’s philosophy to indicate a life force which he conceived of as psychoid or ‘mind-like’, that is; non-spatial, intensive, and qualitative rather than spatial, extensive, and quantitative. The scientific community criticised his concept of entelegy.
66
In Section Adaptation in the 1953 version of Geobiology Baas Becking described “the question of hereditary adaptation” of organisms “in order to make it fit in a new environment” (p. 492-498):
In practice, however, the issue is clouded by the evaluations of the impact of the two agents, heredity and environment, “nature et nourriture”, Blut und Boden”, upon the human race; ecclesiastical. creed demanded a voice, especially in the latter part of the nineteenth century and at present we witness the influence of political creed upon the development of what should be a purely scientific matter. The emotionalism connected with any discussion of this problem has coloured, and thus disqualified, some or the recent work. There is no need to comment upon the great importance of Darwinistic evolutionary thought, especially of the selection principle, as a guide for our understanding of the more passive forces in evolution, for evolution requires a continuous re-adaptation to environment.
The main difficulty seems to be what forces cause this evolution. Is it primarily an inner urge, an inner inevitability, which causes the evolutionary stream to flow and to follow a certain course? Or is this stream directed by the topography the environment? In Section 2 some arguments are given which seem to favour the latter alternative. (See also Umbgrove 1942), In certain geological periods, with sudden changes in climate, periods of orogenesis, sudden veritable explosions of speciation did occur. Was this only the stimulation of the sluggish stream of evolution or the chance of this stream into a new course? And, apart from these evolutionary speculations, is it possible to assume that the “Milieu Exterieur” can permanently modify the hereditary complex?
It has been repeatedly claimed that this is the case. The stimuli may be traumatic, a chemical or physical nature, apart from the (vague) impact of climate. The evidence for mutagenic influence of traumatic stimuli is very doubtful (Sirks, 1951) Climatological influences have been chiefly established by means of populations instead of with genetically homogeneous material. Here again, convincing evidence seems to be lacking. There is no doubt that a substance like colchicine is able to double or to multiply the number of chromosomes in a nucleus, with the formation of a new genotype. The application of high and low temperatures may also cause a change in chromosome numbers. The influence of radium on the genotype has been conclusively shown for plants.
The work on the influence of X-rays, initiated by Muller in 1928 (Drosophila) has shown that radiation is of great influence on the genotype. It was shown that the largest part of the induced mutations was lethal, but that there is a qualitative analogy between he induced and the natural mutations. Later work has shown that the X-rays are effective on plants. Here temperature shocks and ultraviolet light also induce a high mutation rate in the offspring. Cosmic radiation is also effective, both on the mutation rate in fungi and in Drosophila.
Most of the work performed in this field seems to show only that the mutation rate is increased beyond its natural magnitude by certain external influences (Sirks, 1951). The environmental influence would therefore only stimulate the slow stream of mutations which remain autogenous in nature. The induced morphological changes are, on the whole, unspectacular, and not to be compared to specific (let alone generic) differences met in evolution. We are still waiting for a real clue, and we may be extinct as a race before we can solve this Lamarckian-Darwinistic riddle.
67
Reference to Kluyver and Baars (1932). According to Postgate (1979, p. 5-7):
Sulphate reducing bacteria include both ordinary mesophilic strains and thermophilic strains able to grow at temperatures between 50 and 70 °C. Kluyver and Baars (1932) believed that these were adaptive variants of the same organism [..] However, Campbell, Frank and Hall (1956) showed conclusively that the thermophilic types were a completely different species, hitherto known as Closteridium nigrificans […]. Today, the adaptive interconversion of mesophilic and thermophilic species of sulphate reducing bacteria must be regarded as mistaken.
See also Postgate (1995, p. 16-17) and Geobiology, Baas Becking (1953, p. 494):
According to definition of the milieu, the counter mould of life, the sum total or rather, the integration of environmental factors, we will call the “milieu”, every organism adapted to its milieu would be an artifact -, in a physiological, as well as in a morphological sense.
68
Reference to Vaas (1938). See also p. 492-494 in Baas Becking Geobiology (1953), Chapter V, Section Adaptation:
Vaas remarks that the terminology, used for higher organisms; genotype, Phenotype and mutation, is not applicable to bacteria as, without a genome, there will be no genotype. He has coined new terms, hypartype and doxatype while he suggests, the neutral term “pedema” (jump) instead of mutation.
69
In manuscript Geobiology (1953, p. 495), Baas Becking remarked:
It is true that the “pedemata” in bacteria (e.g., white form of prodigious) are often reversible, but so are certain mutations in sexual organs. Obviously, in a nuclear organism the question of hereditary adaptation cannot be settled before sexual reproduction has been convincingly shown to exist. […] But the present evidence of such “bacterial mutations” seems unconvincing.
70
Reference to Otto Rahn (1881-1957), German-American microbiologist and R.E. Buchanan (1883-1973), American bacteriologist and Ellis Ingham Fulmer (1891-1953), American biophysical chemist Iowa State. Fulmer and Buchanan, professors at Iowa State College, wrote the three volume treatise Physiology and Biochemistry of Bacteria (1928-1930). Volume I (1928) dealt with growth phases.
71
Reference to Hof (1935a and 1935b).
72
Johannes Paulus Lotsy (1867-1931), Dutch botanist, Professor of Botany, Leiden University (1904-1909) director Rijksherbarium Leiden (1906-1909), secretary Dutch Society of Sciences Haarlem (1909-1931). Zevenhuizen (2008), gave a detailed description of Lotsy’s controversy with Hugo de Vries (p. 362-370) and Lotsy’s doubts on De Vries mutation theory (p. 375-376).
73
Gyrinus natator, whirligig beetle. Their compound eyes are remarkable for each being divided into a higher part that is above water level when a beetle is floating passively, and a lower part that is below water level. See also Blagodatski. et al. (2014).
74
The reference to Julius Georg Hubertus Wilhelm Troll (1897-1978) is wrong. The term “Fremddienliche Zweckmäßigkeit” was coined by Becher (1917), Die fremddienliche Zweckmäßigkeit der Pfanzengallen und die Hypothese eines überindividuellen Seelischen.
75
Unidentified reference, not found in the 1925 issues of Science or in the bibliographies of Baas Becking and Annand. Baas Becking probably referred to Percy Nicol Annand (1898-1950) an entomologist, who wrote a M.A. thesis in the Department of Biological Sciences at Stanford University in 1922. In 1928 he received a Ph.D. in zoology and botany from Stanford. He described the gall inducing species of the family Adelgidae (Annand, 1928). In 1941 he was appointed as Chief of the US Bureau of Entomology and Plant Quarantaine.
76
Reference to Beijerinck (1897).
For a review of Beijerinck’s studies on galls see van Iterson (1940).
77
See for Kammerer, van Alphen and Arntzen (2016).
78
Reference to Cornelis Hendrik Edelman (1903-1964), Dutch professor in Mineralogy, Petrology, Geology and Agrogeology in Wageningen. Baas Becking probably referred to Edelman (1936). Edelman described the term “rare” as “finely divided material dispersed in small quantities.” Apparently, the Section on Minimum Elements in the present version of Geobiology were strongly inspired by Edelman’s article.
79
‘Locoweed’ is a poisonous plant problem, in the case of Astragalus crotalaria due to Se hyperaccumulating.
80
Baas Becking possibly referred to E. Brandenburg who showed in Holland that crown rot of sugar beet is caused by boron deficiency, and can be cured by the addition of this element. See Brandenburg (1931), Brandenburg (1939).
81
The Cuban soil is high in lithium and thus the cigars have lithium in them too. Hans Goldschmidt in his Erinnerungen an Robert Wilhelm Bunsen (1911), tells the anecdote that as a student he was allowed to make spectral analytic experiments in Bunsen’s laboratory.
As Bunsen was a smoker of strong Havana’s he never failed to point out that in every Havana you could clearly see the red lithium line. If, by chance, a student himself had a cigar which showed no lithium reaction, Bunsen can scarcely ever have missed the opportunity of making a little joke, by pointing out that the student’s cigar was no Havana.
Robert Wilhelm Bunsen (1811-1899), German chemist, he investigated emission spectra of heated elements and discovered caesium (1860) and rubidium (1861).
82
Reference to Richard V. Young, Chemical Laboratory of Iowa State College, who published in the 1930s with Henry Gilman several papers on lithium in the Journal of the American Chemical Society. Baas Becking’s reference to Young (1935) was not identified.
83
According to Gellhorn (1929):
“Die Ionen der Alkalimetalle werden stärker aufgenommen als die der alkalischen Erden. Die Kationen bilden dabei eine Reihe: K > Na > Li > Mg > Ca, und die Anionen folgende Reihe: CnS > Br > J > NO3 > CI > SO4.”
84
Reference to Anneliese Niethammer (1937).
85
Reference to Klement (1938).
86
For Edelman see above. Baas Becking had his reference to Longobardi (1935) from Edelman (1936).
The fungus Amanita muscaria concentrates high levels of amavadine, a natural vanadium complex without the V = O bond.
Henri ter Meulen (1871-1942), since 1905 Professor in Analytical Chemistry, Technical University. Delft. Ter Meulen (1931) found a remarkable accumulation of vanadium (120 ppm dry weight) in A. muscaria.
87
Baas Becking referred to information on vanadium from J.M. Boutwell in Utah in Clarke, 1916 (p. 716-717 in the 1920 fourth edition of Clarke’s Data of Geochemistry).
88
Manganese is an essential plant micronutrient. Mn toxicity and deficiency in plants are documented. In practice there is a delicate balance between the microbial oxidation of Mn2+ and the reduction of Mn oxides, which can be shifted by small changes in soil water potential along with changes in water temperature and pH. See Sparrow and Uren (2014).
89
Baas Becking also referred to Leo Minder in Section 8.5.5.
90
Reference to Beger (1935), Leptothrix Echinata, ein Neues Vorwiegend Mangan fällendes Eisenbakterium. Leptothrix is an aerobic chemolitotrophic bacterium that uses Mn2+ for Manganese oxidation (MnO2).
91
Reference to the work of H.O. Askew and H.R. Marston on ‘Coast disease’ of sheep in New Zealand and South Australia. Due to cobalt deficiency. See for a review and literature references; Beeson (1950).
92
Reference to Raulin (1869). Raulin demonstrated the essentiality of salts of Zn, Fe and Mn as well as silicates for the growth of Aspergillus niger. We found no reference to the stimulating working of arsenic.
93
See Kudo (1954), Protozoology. Baas Becking doubtlessly used an early edition.
94
Reference to Otto Bütschli (1848-1920).
95
Ter Meulen (1931) succeeded in separating and estimating the molybdenum present in various plants and animals. See also H. ter Meulen (1932), Distribution of Molybdenum. Nature 130, 966.
Bortels (1930). Bortels first established the importance of molybdenum and vanadium for nitrogen fixation, and also noted, albeit only in a footnote, that Azotobacter vinelandii could grow in the absence of both Mo and V. See Pau (1993).
96
The extraction of iodine from Chilean nitrate was before WWII the main source of iodine. The main difference between sodium iodide and sodium chloride is that sodium chloride is composed of a chloride ion bonded to a sodium ion whereas sodium iodide is composed of an iodide ion bonded to a sodium ion. Sodium iodate can be oxidised to sodium periodate by strong oxidising agents: NaIO3+NaOCl →NaIO4+NaCl.
97
Goitre is swelling in the neck resulting from an enlarged thyroid gland. Over 90 % of goitre cases are caused by iodine deficiency.
98
Mulder (1938). Mulder used in his tests the colour of the spores of Aspergillus niger grown in a copper deficient medium supplied with small amounts of soil as a measure of Cu supply to the fungus. This served as a measure for the availability of Cu to cereal plants growing on newly reclaimed peaty and heath soils.
99
Baas Becking referred to a personal communication of H.G. Derx.
100
Reference to Jan Smit (1885-1957), Professor of Microbiology, Wageningen (1936-1956). The reference is to Smit (1930) and possibly also to Smit (1933).
101
V.M. Vernadsky was with Lotka (1924) and Henderson (1913) one of Baas Becking’s main inspirators for Geobiology. Here he referred to the first part of La Géochimie (1924). In the 1998 English version, The Biosphere (translation of the original edition in 1925 in Prague), the vital explosions are treated in the paragraphs 25-33.
102
In the 1953 version of Geobiology, Section on Man in Relation to Pathogenic Organisms (p. 701), Baas Becking included a reference to:
Hans Zinsser in his remarkable book on Rats, Lice and History (1935) has sufficiently stressed the importance of certain pathogens in human history, which is still taught as a story of battles, treaties and conventions, although the factors of plague, malaria, spotted typhus and tuberculosis have been more potent than many a human plan.
Human history has been punctuated by epidemics. Homer mentions a plague epidemic during the Trojan war, and the black death of 1346, which killed at least three quarters of the population of Europe is still remembered. Epidemics have continued in the intervening centuries, and the last fifty years can show most striking examples as the outbreaks of yellow fever at Panama, of dengue at Athens and of typhus in the Balkans. Human health and population growth have been greatly affected, both quantitatively and qualitatively by the recognition of pathogenic agents and by the means devised for their control.
103
Reference to Dr. G.P.H. van Heusden, biologist of the Amsterdamse Waterleiding. He obtained his Utrecht PhD in 1943 on a study of the migration of glass eel to Lake IJssel: De Trek van den Glasaal naar het IJsselmeer.
104
See for Hugo de Vries’ research in Rotterdam, Zevenhuizen (2008, p. 142-143, and note 99 on p. 554).
105
Reference to Baas Becking, Tolman, McMillin, Field and Hashimoto (1927). In the 1953 manuscript Geobiology on p. 526-528 under the heading Silica, Baas Becking described the diatoms research in more detail.
106
In his Inaugural Address in Utrecht University Baas Becking (1927) referred to Vernadsky’s “explosions de la vie” and referred to the Aulacodiscus kittoni epidemy and Forti’s ‘Mare sporco.’
Masses of a yellowish or greenish-brown gelatinous substance floating at the surface of the sea and at times colouring large areas are the cause of the phenomenon known as ‘mare sporco.’ It is especially noticeable in the Adriatic, but may occur also in many other places. It seems that these curious masses in the water have their origin at the bottom where innumerable diatoms, usually a few definite bottom living species, in certain circumstances by rapid reproduction give rise to a large amount of a gelatinous substance impregnated with much oxygen.
Forti (1906). See also The Origin of ‘Mare Sporco, Nature (1932, 129, p. 660).
107
Pelagia noctoluca are bioluminiscent. Light is emitted in the form of flashes when the medusa is stimulated by turbulence created by waves or by ship’s motion. In his Inaugural Address in Utrecht University (Baas Becking, 1927) referred to a personal observation in the Bay of Tehnantepec, that in front of the ship there was during more than two hours a mass of the light emitting medusae, he estimated their biomass as several million tons.
108
Carl Johan Joseph Ernst Wiman (1867-1944), Swedish palaeontologist. Reference not identified.
109
Pachychilus is the generic name of the jute snails an aquatic gastropod. Baas Becking probably referred to Locusta migratoria. The species name tartarea is obsolete.
110
More correct is I Samuel 6: 1-21.
111
Baas Becking referred to the Wieringermeerpolder, newly created land, developed in the 20th century by draining parts of the inland Zuiderzee. See also notes Sections 7.9.11 and 8.5 and Feekes (1936).
112
Baas Becking probably referred to Vanessa cardui (Lepidoptera Nymphalidae), Pieris spp (Lepidoptera Pieridae) and the western tiger swallowtail Papiliorutulus.
113
The reference is to Baas Becking (1931c).
114
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. In the 1953 version of Geobiology (p. 124) he referred, this time not only with a reference to Nicolle’s Naissance, but also with a quote. The reference to Nicolle is preceded by a personal note on Baas Becking’s experience in the Siegburg prisoner camp in February-May 1945:
Symbiotic relations between two and more organisms, extensively studied in epidemiology (e.g., Ross’ malaria equations) often give rise to a temporary preponderance of one or two components. The author has witnessed in a German prison camp (January-May 1945) where the explosive nature of the Rickettsia infection was clearly indicated (B.B., 1945). Several months before the outbreak the infestation with body lice was almost complete. […]. The onset of the disease was but little steeper than its decline. The epidemic disappeared in the spring apparently more or less independent of the modern hygiene and prophylactic measures of our American liberators!
Nicolle, a Nobel Laureate (because of his fundamental work on typhus epidemics) has described the explosive nature of such calamities in his classic Naissance, Vie et Mort des Maladies Infectieuses. He states (p. 82) “lorsque les propriétés virulents de certains microbes pathogènes sont portées à un point extrême en que la contamination se trouve favorisée par de grandes facilites de contact, mieux encore si ces deux conditions se rencontrent à la fois, les maladies peuvent frapper dans un temps très court un grand nombre d’individus appartenant à la même espèce. Il y a alors épidémie.”
The social implications of such an outbreak are vividly portrayed by Camus (1947) [La Peste] in his description of a plague epidemic in Africa. The quote is on p. 82-83 in Charles Nicolle (1930), Naissance, Vie et Mort des Maladies Infectieuses.
The reference to ‘BB, 1945’ is to L.G.M. Baas Becking, editor. The Typhus Epidemic at Siegburg Penitentiary in 1945. Siegburg April/May 1945. Baas Becking wrote in the period April 10, 1944 until May 23, 1944 a report about the typhus epidemic in the Siegburg prisoner camp during the time he and the former prisoners were in quarantine. There is also a report of the German physician Jacob Ahles about the epidemics written in 1946 in German. Ahles (1946), Die Fleckfieberepidemie im Zuchthaus Siegburg im ersten Halbjahr 1945. March 1946. See NIOD 250c, nr. 167; NIOD 250c, nr. 57.
115
Baas Becking referred to Chapter VIII of A. Lotka (1924), where several examples of equations used in the epidemiology of malaria and parasitology are discussed on p. 79-99.
116
This section on mass death consists of short notes on sources that have not been developed by Baas Becking into a text.
117
Apparently references from a 1929 copy of Natur Und Museum from the Senckenberg Institute. Baas Becking possibly had a copy in the Utrecht prison.
118
Wilting coefficient is the level of soil moisture at which water becomes unavailable to plants and permanent wilting ensues.
119
Dr. J. Zonneveld, not identified. He must not be confused with the Wageningen Professor in Landscape Ecology Isaak Samuel (‘Ies’) Zonneveld (1924-2017), who started his research of the soil and vegetation in the Biesbosch tidal area in 1943 during WW II.
120
Reference to Walter (1931).
121
Baas Becking referred to MacDougal (1924) and to Peirce (1901).
For fog interception by mature coast redwood Sequoia sempervirens trees see Simonin, Santiago and Dawson (2009).
122
Reference to Wood (1925).
123
For Heinrich Walter see Sections 1.2 and 5.10.4.
124
Baas Becking referred to the dissertation of A. Quispel (1943, 1946) in which the findings of K.I. von Goebel on the water relations of the lichen thallus were discussed. According to Goebel the fungus protects the alga against desiccation by its water reserve in the “Quellhyphen.” It appeared from Quispel’s experiments (p. 522):
[…] that this protection against desiccation could only be observed at a low degree of desiccation of the thallus. On the contrary it appeared that in consequence of the symbiosis the algae are much more dependent of liquid water than had been the case in a free living state. The presence of this water reservoir in the hyphae will be only of minor importance for the gonidia and certainly the benefit will be much too insignificant to base a mutualistic theory of the symbiosis upon.
See Goebel (1926a), Goebel (1926b), Goebel (1927).
125
Baas Becking probably means the reduction potential of oxidising agent and reducing agent, which has the reduction potential V = 0.
126
The reference to Suringar about ‘budding’ not identified. Baas Becking referred to Willem Frederik Reiner Suringar (1832-1898), Dutch botanist, Professor of Botany and director Hortus Botanicus Leiden University (1862-1898) and director Rijksherbarium Leiden (1871-1898). His son was Jan Valckenier Suringar (1864-1932), from 1900 till his death in 1932 director Wageningen Botanical Garden. Baas Becking’s cousin Louise Hermine Baas Becking (1881-1969) was assistant of Valckenier Suringar until her marriage with F.A. Moraux in 1921. She was responsible for the design of the Wageningen Botanical Garden De Dreijen (‘Baas Beckingtuin’).
Baas Becking wrote a well informed outline of the scientific work of W.F.R. Suringar, his predecessor in Leiden. See Veendorp and Baas Becking (1938, p.170-177).
127
According to Hecht (1933) the influence of H2S, a product of decomposition of organisms, on the benthos has been overestimated. He found that Arenicola and Nereis were not affected by a temporary content of hydrogen sulphide in the water. He also called attention to the fact that, even if hydrogen sulphide is found in the sediment, as is indicated in fossil sediments by the occurrence of pyrite, we cannot definitely conclude that it was present in the water, and only the quantity present in the water influences the benthos.
128
Barcroft and Barcroft (1924, p. 41-42) owed that the red blood of Arenicola contains a form of haemoglobin different from that contained in human blood. The oxygen dissociation curve shows a higher affinity for oxygen, so that the gaseous exchange takes place at very low pressures. The blood volume of Arenicola varies greatly in different worms, but the oxygen capacity is much more constant, being of the order of 0.01 cc of O2per g of worm. This quantity is definitely less than the maximum the Arenicola is estimated to need during the time when its hole is sealed up at low water. Nevertheless, it is so nearly of the same order as to make clear the probable function of the pigment, namely, to act as a reserve store which the organism can use up when it has not access to seawater.
129
Quote from Barcroft and Barcroft (1924, p. 36).
130
Reference to Maurits Henri van Raalte (1907-2002), plant physiologist at Treub Laboratory Buitenzorg (1939-1956), later Professor of Plant Physiology, Groningen University. See van Raalte (1941).
131
In the 1953 manuscript of Geobiology (Baas Becking, 1953a, p. 492) Baas Becking referred to:
Brongersma-Sanders (1943) has called attention to the mass death of fish and molluscs, which occurs periodically in an area of Walfish Bay, South Africa. Here sediments rich in FeS and in organic matter are found which are apparently, apart from bacteria, deprived of life. She ascribes this mass death to a water bloom of a Dinoflagellate. These organisms are reputedly toxic to fish and are the cause of similar calamities in other regions of the world. Sulphate reduction and the subsequent exhaustion of the water in oxygen might be a secondary phenomenon.
132
Reference to the Snellius expedition, an oceanographic survey of the waters of eastern Indonesia from July 1929 until November 1930. H.J. Hardon, Ph. Kuenen and H. Bosma were participants of the expedition, the scientific results were published by Brill. The sea voyage was led by lieutenant F. Pinke, brother of Baas Becking’s sister in law Eliza Hendrika Pinke (1898-1942).
‘Ocean sill’ is a sea floor barrier of relatively shallow depth restricting water movements between ocean basins.