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TitleBeyond the Gene - Cytoplasmic Inheritance and the Struggle for Authority in Genetics - J. Sapp (Oxford, 1987) WW
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Beyond the Gene

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with a Nobel Prize (see Chapter 7). It is little known, however, that Ephrussi also
played a leading role in the institutional development of French genetics and in
fostering research on cytoplasmic inheritance and challenging the predominant
role of nuclear genes in heredity and evolution.

Immediately following World War II, Ephrussi was appointed to the first chair
of genetics at the Sorbonne and became largely responsible for the development
of genetics in the French university curriculum. After establishing an institute for
genetic research in France in 1946, Ephrussi organized a comprehensive research
program which centered on investigations of genetic regulation based on cyto-
plasmic inheritance in Drosophila, Podospora, and yeast. The research carried
out by Ephrussi and his co-workers on yeast concerned the non-Mendelian in-
heritance of respiratory-deficient petite mutations. Continued investigation of this
characteristic for more than a decade would ultimately be regarded as providing
the first genetic evidence for mitochondrial heredity (see Chapter 7).

The Neo-Lamarckian Hegemony

The Napoleonic structure was rigidly hierarchical. It was a mixture of an eccle-
siastical control of ideas, governmental bureaucracy, and the military style of the
emperor. (Terrence Clark, 1973, p. 18)

Ephrussi was born in a suburb of Moscow and was initially introduced to genetics
as a university student. Following the Bolshevik Revolution, he left Russia for
Rumania, where he tried to follow the Tolstoyan way as a farmer. After a year
and a half of this way of life, Ephrussi disavowed himself from agriculture and
religion and emigrated to France to return to science at the University of Paris.
Though genetics had established itself in Britain and the United States by the
1920s, it had been resisted in France. In direct conflict with the United States and
England, classical Mendelian genetics was almost nonexistent in French univer-
sities, which remained a bastion of neo-Lamarckism (Boesiger, 1980; Limoges,
1980; Buican, 1984).

Neo-Lamarckism had emerged in France during the 1880s when Darwinian the-
ory, which had previously included the notion of the inheritance of acquired char-
acteristics, was beginning to be identified solely with natural selection (see Chap-
ter 1). Since that time almost all biologists in France not only acknowledged a
role for the inheritance of acquired characteristics but used it as a central deter-
minant in an explicitly anti-Darwinian view of evolution. A nationalistic com-
ponent contributed to the emergence of the neo-Lamarckian viewpoint in France.
The defeat of France in the 1870 war with Bismarck's Prussia brought with it a
patriotic tendency of French biologists to encourage support of their own theories
rather than to accept or develop those of other countries, especially Germany. In
the face of the impending neo-Darwinian views, French biologists upheld La-
marck, attributing to him notions that were quite alien to those conveyed in his
writings (Boesiger, 1980; Limoges, 1980).

World War I helped to further imbed Lamarckism in French patriotism. In fact,

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Lamarckism in France came to represent a form of nationalism which was posited
in direct opposition to Germany and German militarism with Bismarck. Traces
of this anti-German sentiment can be detected in much of the French biological
literature between the two World Wars. When Maurice Caullery, who held the
chair of evolutionary biology at the Sorbonne, evaluated the contributions that
diverse countries had made to the development of biology, he wrote:

It is necessary after having escaped the peril of Germanic political hegemony, not
to lose sight of another danger which threatens us, in the name of unjustified pre-
tensions, that of the intellectual hegemony of Germany. (Caullery, 1922, p. 23, my

It is worth mentioning in passing that German biological descriptions of cellular
processes were not without their military metaphors of machine guns and armies.
In 1885, the leading neo-Darwinian Weismann (p. 195) described the action of
the nucleus during development in the following metaphorical terms:

The development of the nucleo-plasm during ontogeny may be to some extent com-
pared to an army composed of corps, which are made up of divisions, and these of
brigades, and so on. The whole army may be taken to represent the nucleoplasm of
the germ-cell: the earliest cell-division . . . may be represented by the separation
of the two corps, similarly formed but with different duties; and the following cell-
divisions by the successive detachment of divisions, brigades, regiments, battalions,
companies, etc.; and as the groups became simpler so does their sphere of action
become limited.

The Mendelian-chromosome theory as formulated by the Morgan school stood
in virtual conflict with some of the central presuppositions of the neo-Lamarckian
approach to heredity. Neo-Lamarckian biologists such as Yves Delage viewed
heredity as an epigenetic process (Fischer, 1979). The adult characteristics of the
organism were seen to result from a series of complex and integrated processes.
Both intraorganismic and extraorganismic environmental circumstances contrib-
uted to this process. As discussed in Chapter 1, from the epigenetic perspective
of the organism as a whole, any particulate theory of "determiners" seemed to
be wrong and naive.

For Delage (1903, p. 806) the germ plasm of the egg contained only two es-
sential factors, a relatively simple chemical composition and an arrangement of
its parts. Delage searched for the causes of variations and their transmissibility in
problems of ontogenesis. Lamarckism was a "somationist" theory of evolution
which presupposed that characteristics acquired by the soma could have a specific
influence on the germ. There could be no fundamental separation of the somato-
plasm from the germ plasm. For example, Delage thought that an amputation of
a gland could have a correlative effect and become hereditary. He attached great
importance to nutrition, which he considered to have a morphogenetic action by
adding to the chemical composition of cells, and modifying the nature of the
substances and their arrangement. In his view, the Irish, English, and Arabs ob-
tained their "racial" characteristics from differences in their diets (Delage, 1903,
p. 836).

The views of Delage were representative of the thinking of many French bi-

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Sorbonne, 127, 181, 188, 189-91
Soviet Academy of Science, 172
Spatial principle. See Polarity
Species characters, 16, 54, 56, 74, 99, 126. See

also Fundamental characters
Species specificity, 21, 23, 29
Spemann, H., 14, 24, 60, 67-70
Spencer, H., 5
Spiegelmann, S., 102, 103, 115, 136-37, 154
Springarn Medal, 143
Stahlian theory, 41
Stalin, J., 165
Stanford University, 201, 205
Statistics, and genetics, xii, xiii, 33, 36, 39-40,

60, 93. See also Biometry; Population genetics
Steady states, 101, 151, 168, 196, 198, 199-202,

Stent, G., 211
Stern, C., 57
Strasburger, E., 4
Structural guidance, 218
Structural inertia, 218
Stubbe, H., 73
Sturtevant, A. H., on cytoplasmic inheritance,

27-28, 82, 100-101, 104, 148-49, 209
Substratum, 66
Sumner, F. B., 45-46, 224
Superficial characters, 22, 16, 17. See Funda-

mental characters
Supergene, 19, 20. See also Vitalism
Supramolecular structure, 92, 93, 109, 212-20,

Sydney Farber Cancer Institute, 210
Symbionts, xii, 84. See also Endosymbiosis;

Symbiosis, 10, 18. See also Endosymbiosis
Symmetry, 9, 12, 13, 14, 21, 27-28. See also


Tartar, V., 216
Tatum, E. L., 98, 120, 123
Technique-ladenness of observations, xv, 25, 28,

156-58, 226-27
Teissier, G., 128-29, 134, 140, 182, 184-87
Ternitz, C., 79, 169
Tetrahymena genetics, 197, 217
Totipotency, 6, 33
Tower, W. L., 92
Toyama, K., 27
Transductions, in bacteria, 121
Transformations, in bacteria, 121, 169
Tschermak. E. von, xii
Tubingen, 75
Typological thought, 84-85

United States Department of Agriculture, 55
University of Berlin, 57
University of Brussels, 145
University of California, Berkeley, 66
University of Chicago, 143
University of Cincinnati, 17
University College, London, 170
University of Edinburgh, 105
University of Glasgow, 153, 189
University of Missouri, 205
University of Paris, 127, 128, 190. See also Sor-

University of Sussex, 220
University of Wisconsin, 62-65

Vavilov, N. I., 163
Vernalization, 164
Viruses, 89, 102, 117-22, 214
Vitalism, 7, 19-20, 69, 91, 126

Waddington, C., 99, 166-67
Wagtendunk, W. von, 120
Wallace, A. R., 4
Wallin, J. E., 121
Washington University in St. Louis, 102, 154
Watson, J. D., xiii, 114, 189, 193
Weaver, W., 110, 111, 114-15, 154, 181-88.

See also Rockefeller Foundation
Weidenreich, F., 57
Weindling, P., 58
Weismann, A., 4-6, 125, 222
Weismannism, 7, 40, 41, 48-50, 125, 212
Weiss, P., 8, 14, 69, 143
Weldon, W. F. R., 33, 36
Weltanschauung, 68
Went, F., 176
Wersky, G., 166
Wettstein, D. von, 74, 75
Wettstein, F. von, 56, 73-77
Wettstein, R. von, 57-58
Wheeler, W. M., 51
Whitman, C. O., 7-11, 18
Wiggles worth, V. B., 152
Wilson, E. B.: on cytoplasmic particles, 25-26;

earlier views on exclusive nuclear heredity, 4,
6, 10-12, 117; on ground substance, 26; and
Morgan school, 45; on organism as a whole,
10; on organization of the cell, 34

Winge, O., 136
Winkler, H., 72, 75, 76
Woolf, V., xi
Wright, S.: on cellular differentiation, 99, 100-

101; on Dauermodifikationen, 101; Plasma-
gene theory of, 100, 106, 147; and population

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Wright, S. (continued) Yeast genetics, 102-4, 133-36, 146-51, 154-
genetics, 63, 85, 93; steady state, concept of, 55, 158, 170, 202-3, 208-9
101, 196 Yoxen, E. J., 154

X-rays, 51—52, 62, 93, 113 Zeitschrift fur Induktive Abstammungs und Ver-
erbungslehre, 57

Yale University, 205 Zirkle, C., 173

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