Physiology and the whole organism  ·  Article

Chemical control of the sign and precision of heliotropic response

Loeb reasoned that if the immediate effect of light in causing heliotropic reactions is chemical, chemicals should be able to control both the precision and the sense of the response.1 He tested this on the fresh-water crustaceans Gammarus, Daphnia and Cyclops.2 Acids, above all carbon dioxide, turned indifferent or negative animals toward the light.4 Loeb noted that it may be of importance that these chemicals are ones the organism itself may form.1

Earliest held
1890, James, W.
Most discussed in
The Cell in Development and Inheritance, 1896
In the library
107 passages in 13 works
Rewritten
2026-11-07
01

Gammarus and the acid test

In 1906 Loeb reported that Gammarus left alone cling to solid bodies anywhere in the aquarium. If disturbed, by being moved or by stirring the water, they become negatively heliotropic, at least for a time. Adding certain chemicals such as esters made them at once intensely positively heliotropic.3 He could not obtain this alkali effect in Gammarus.4

02

Cyclops, Daphnia and temperature

In Cyclops, adding HCl, another not too weak acid, or carbon dioxide made the animals all gather in a narrow region at the window side of the vessel.4 Loeb summed up the Cyclops result by saying that acid, especially carbon dioxide, made negative Cyclops positive. It also strengthened the positive response of animals that were already positive.4 In 1912 he wrote that Daphnia usually need the water cooled as well. Cooling plus carbon dioxide made light-indifferent Daphnia strikingly positive.5 He also stated that marine copepods can be made positive by lowering the temperature alone, or by a sudden rise in the concentration of the sea water.5 In 1918 he restated the finding for Californian Daphnia, copepods and Gammarus, using weak acid, especially carbon dioxide.6

03

Colour and intensity disputes

Other authors, as summarised by Washburn in 1908, studied Daphnia's response to light without chemicals. Von Frisch found that a blue screen made Daphnias avoid light even though it lowered the intensity.7 Ewald reached conclusions about colour blindness in the side regions of the eye. Washburn called this a heavy weight of inference on slender evidence.8 Loeb and Moore reported that ultra-violet rays instantly made Daphnia avoid the light containing them. Lubbock found that Daphnia sought such light in preference to darkness.8 These reports suggest that the sign of the response depends on more than intensity, though the evidence is mixed. Washburn doubts Ewald's inference and notes that Von Frisch is determined to find colour vision in invertebrates. Lubbock's finding also contradicts that of Loeb and Moore.7

04

A related salt effect

In 1916 Loeb wrote that the results of his Fundulus experiments are of general application. Osterhout had shown that plants of soil or fresh water are readily killed by pure NaCl at a certain concentration. They survive the same concentration if some CaCl2 is added.9 Ostwald had shown the same for a species of Daphnia.9 These results concern toxicity, not orientation to light.

SourcesEach quotation was checked word for word against the passage it opens.
  1. we should expect that it must be possible by the use of chemicals to control the precision and sense of the heliotropic reactions.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 353
  2. The experiments were made on fresh-water Crustaceans, Gammams pulex, Daphnia, and Cyclops.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 353
  3. It is possible, however, to make them at once intensely positively heliotropic, by adding certain chemicals to the water, e.g. esters.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 353
  4. We see, therefore, that acid, especially CO2, not only makes negatively heliotropic Cyclops positively heliotropic, but increases the intensity of the positive heliotropism in those that were already positive at the beginning of the experiment.Loeb, J., 1906 · The Dynamics of Living Matter · open at passage 357
  5. But if a blue screen is interposed between the light and the animal, in spite of the fact that the intensity is thereby diminished, the Daphnias avoid itWashburn, M. F., 1908 · The Animal Mind: A Textbook of Comparative Psychology · open at passage 251
  6. On the other hand, Loeb (431) and Moore (so02) report that the action of ultra-violet rays instantly makes Daphnia avoid the light containing themWashburn, M. F., 1908 · The Animal Mind: A Textbook of Comparative Psychology · open at passage 252
  7. If the water containing the Daphniae is cooled and at the same time carbon dioxide added, the animals which were before indifferent to light now become most strikingly positively heliotropic.Loeb, J., 1912 · The Mechanistic Conception of Life · open at passage 392
  8. Wo. Ostwald3 has shown the same for a species of Daphnia. We, therefore, come to the conclusion that the injuriousLoeb, J., 1916 · The Organism as a Whole, from a Physicochemical Viewpoint · open at passage 476
  9. certain fresh water crusta¬ ceans, namely Californian species of Daphnia, copepods, and Gammarus when indifferent to light can be made intensely positively heliotropic by adding some acidLoeb, J., 1918 · Forced Movements, Tropisms, and Animal Conduct · open at passage 195
Linked ideas
Covers the orientation of organisms to light and gravity, the response whose sign Loeb altered with chemicals.
Tropismpart of / contains
Heliotropic reversal by acid is one of Loeb's experimental cases for treating orientation as a chemical and physical process.
Gives the photochemical account of heliotropism that the idea of chemical control of its sign and precision builds on.
Osterhout and Ostwald showed that CaCl2 offsets NaCl toxicity, including in Daphnia, another case of chemicals shifting organism response.
Carbon dioxide and acids acted as the main chemical switches in these experiments, linking them to acid-base chemistry.
Loeb restated the acid results in his 1918 book on forced movements, tropisms and animal conduct.
Where it is discussedPassages matching control of heliotropic reactions by chemicals, Gammarus, Daphnia, Cyclops
1896The Cell in Development and Inheritance · Wilson, E. B.27
1906The Dynamics of Living Matter · Loeb, J.20
1908The Animal Mind: A Textbook of Comparative Psychology · Washburn, M. F.16
1918Forced Movements, Tropisms, and Animal Conduct · Loeb, J.15
1912The Mechanistic Conception of Life · Loeb, J.9
1916The Organism as a Whole, from a Physicochemical Viewpoint · Loeb, J.7
1915Principles of General Physiology · Bayliss, W. M.4
1930The History of Biological Theories · Rádl, E.2
1890The Principles of Psychology, Vols. 1-2 · James, W.2
1894Materials for the Study of Variation Treated with Especial Regard to… · Bateson, W.2