Bowler - Theodor Eimer and Orthogenesis

Theodor Eimer and Orthogenesis:
Evolution by 'Definitely Directed Variation'

PETER J. BOWLER
I U M E R O U S objections were raised against Darwin's

theory of evolution by natural selection during the late
nineteenth and early twentieth centuries.1 One of the
most interesting objections concerned the relationship
between variation and selection as the factors determin-
ing the course of evolution. Darwin assumed that the
variations of individuals occurred more or less at random, hence selection
for advantageous characters was the main directing agent. But if variations
were not randomif they tended to occur more readily in some directions
than othersthen the direction of variation might itself control the course
of evolution, with selection playing only the negative role of eliminating
those directions that were harmful. 'Orthogenesis' was the name coined
by Wilhelm Haacke and popularized by Theodor Eimer to designate the
process of evolution by 'definitely directed variation.'2 The name ortho-
genesis soon became popular, especially among the paleontologists. It is
still associated largely with those trends observed in the fossil record that
seem difficult to explain by natural selection.3 However, Eimer himself
supported orthogenesis with studies of living species, and the late nine-
teenth century saw an extensive debate over the value of this kind of
evidence, a debate which seems worthy of the historian's attention. Eimer
and his followers knew of the fossil evidence for orthogenesis, but felt that
1. On the background to the anti-Darwinian movement see, for instance, Loren Eiseley, Darwin's
century: evolution and the men who discovered it (New York, 1958), chapter 9; and Philip G. Fothergill,
Historical aspects oforganic evolution (London, 1952). A useful contemporary account is Vernon L.
Kellogg, Darwinism today: a discussion of the present-day scientific criticism of the Darwinian selection theo-
ries . . . (New York, 1907).
2. Gustav Heinrich Theodor Eimer, On orthogenesis and the impotence of natural selection in species
formation, trani. J. M. McCormack (Chicago, 1898), p. 19. Eimer mentions Haacke's prior use of the
term in Wilhelm Haacke, Cestaltung und Vererbung (1893). The term comes from the Greek 6pd6t,
straight and ytvtait, production.
3. See for instance Julian Huxley, Evolution: the modem synthesis (1942; reprint ed.. New York,
1964), pp. 497-525; and George Gaylord Simpson, The meaning of evolution, rev. ed. (New Haven,
1967), chapter 11.
[40]
Bowler : Theodor Eimer and Orthogenesis 41
their own investigations were equally important since they appeared to
provide experimental disproof of Darwin's theory. The opponents of or-
thogenesisthe chief of whom was August Weismanntried to minimize
the significance of the whole affair so as to leave as much as possible of the
original structure of Darwinism intact. Weismann, die originator of the
germ plasm theory, polarized opinions within evolutionism by advocating
an exclusive form of selectionism which became known as neo-Darwin-
ism. His consistent opposition to Lamarckism is well known, but Weis-
mann also felt that orthogenesis presented a significant threat to his theory.
In the end Weismann himself was forced to concede at least some directed
variation, so the debate with Eimer gives a dear indication of the perilous
state of Darwinism at the end of the century.
Eimer's work has received little attention from historians, apart from a
brief account by Nordenskiold.4 Very little biographical information is
available.5 In part this neglect may be due to the fact that ordiogenesis
came to be associated more with trends in the fossil record than with the
kind of evidence advocated by Eimer, but it also stems from the original
debate with Weismann, whose fame bodi then and now has tended to
overshadow Eimer's contributions. Certainly Eimer himself felt that the
machinations of Weismann were responsible for his failure to gain the
international recognition he thought he deserved. His writings became
known for their excessively polemical tone, which seems to have discour-
aged some potential followers.6 Eimer also tended to push his ideas to
extremes that encouraged rejection even by diose who were hostile to
Weismann's neo-Darwinism. Thus the English anatomist Bertram Windle
wrote of the debate between the 'dreamer of Freiburg' (Weismann) and
the 'visionary of Tubingen' (Eimer), concluding that each succeeded only
in destroying the other's position without establishing his own.7 Eimer
was particularly unhappy with the failure of his work to gain access to the
English speaking world, especially when compared to the frequent trans-
lation enjoyed by Weismann. Only one of Eimer's major works was trans-
lated, and it dealt more with Lamarckism than orthogenesis.8 Of all Ei-
4. Eric NordenskiGld, The history of biology (1928; reprint ed., N e w York, 1946), p. 570.
5. Biographical information on Eimer is derived mainly from the obituary by his pupil, the Coun-
tess Maria von linden, Biologisches Centralblatt, 1898, 1S, 721-725.
6. See for instance the reaction of Kellogg, Darwinism today (n. l ) , p. 321.
7. [Bertram Windle], review of Eimer, Organic evolution in Edin. Rev. 1890, 172, 316-349.
8. Theodor Eimer, Organic evolution as the result of the inheritance of acquired characters according to the
laws of organic growth, trans. James T. Cunningham (London, 1890), translation of volume 1 of Die
Entstehung der Arten auf Grund von Vererbung erworbener Eigenschaften nach den Cesetzen organischen
Wachsens (Jena, 1888).
42 Journal of the History of Medicine : January
mer's writings on orthogenesis, the only one translated was a short address
delivered to the Zoological Congress of Leyden in 1895.9 Weismann's
address to the same congress proposed his own mechanism for directing
variation, germinal selection, and was also translated.10 Like the rest of
Weismann's writings it played down the significance of Eimer's theory
and (what incensed Eimer all the more) did so without even discussing
the relevant observations.
Gustav Heinrich Theodor Eimer was born near Zurich in 1843. He be-
gan his studies at Tubingen in 1862, and in 1868 took a medical degree at
Berlin. He also studied zoology, beginning with a short period under
Weismann at Freiburgan interlude about which one would like to know
more than is revealed by the available sources. In 1869 he took a D.PHIL.
degree at Wurzburg. From 1869 to 1874 he appears to have studied inde-
pendently, and made a trip to Italy where he worked on the unusually
colored lizards of Capri. In 1874 he became Inspector of Museums at
Darmstadt, but in the following year moved to the University of Tu-
bingen where he eventually became professor of zoology. He grew in-
creasingly dissatisfied with Darwin's selection theory, turning to both La-
marckism and orthogenesis as alternatives. His principal line of evidence
for orthogenesis was derived from an extensive comparison of various
species of butterflies, but by the end of his life he was speculating on or-
thogenesis as a major factor in all aspects of evolution. He died suddenly
in 1898.
THE ROLE OF VARIATION
Although Darwin had realized that variation is normally random, he knew

that variation was not uncaused and speculated that it was stimulated
chiefly by the environment. Selection was essential because in normal cir-
cumstances variation simply made the species 'plastic,' so that it was flex-
ible to change in all directions. Many different kinds of variation appeared,
and since variation itself was not directed toward any particular goal, selec-
tion was needed to pick out certain factors and eliminate the rest.11 But
from the beginning Darwin admitted exceptions to this rule. Most ob-
vious were the inherited effects of use and disuseLamarck's mechanism
9. Eimer'i address is hii Orthogenesis (n. 3).
10. Wciimann'i address ii Ongeminal selection (Chicago, 1896), alio translated in The monist, 1895-
96, 6, 350-293. Allreference!below are to Ac latter source. Eimer's complaint that Weiimann ig-
nored hii evidence wai not strictly speaking correct, see p. 254.
11. For a typical description of the normally random nature of variation tee Charles Darwin, On
the origin ofspecies by means of natural selection... (London, 1859), pp. 131-132.
Bowler : Theodbr Eimer and Orthogenesis 43
12
of the inheritance of acquired characteristics. In such cases variation was
directed by the habits of the species and was always adaptive. Lamarckism
was sometimes associated with orthogenesis, especially when paleontolo-
gists such as Edward Drinker Cope used the concept to explain linear
trends in the fossil record.13 Cope believed that use-inheritance would
allow the existing habits of a species to direct its evolution over long pe-
riods of time, producing trends that were both linear and adaptive. It was
the consistency with which the variation was supposed to be directed by
habit that allowed the connection with orthogenesis to be made, not the
adaptive goal. In his original definition of orthogenesis, Eimer himself
carefully distinguished between the adaptive trends of Lamarckism and the
direction of orthogenesis which had no utilitarian value. The distinction
was important, since the nonadaptive trends were by far the more difficult
for Darwin's followers to deal with. As early as 1862, Henry Walter Bates
had pointed out that the consistent action of selection might produce trends
that would appear to be the result of directed variation.14 Such an effect
was later given the name 'orthoselection.'15 But an explanation in terms of
selection could only be applied in the case of adaptive trends, while non-
adaptive orthogenesis struck at the basic foundations of the selection theory.
There were two factors which appeared to be capable of producing
directed nonadaptive variation, both of which were mentioned by Dar-
win. One was the direct action of the external conditions which, in some
circumstances, might make all the individuals vary in the same direction.
As an example of the influence of the external conditions Darwin men-
tioned cases where individuals at the edge of a species' geographical range
differed in some (nonadaptive) way from the rest of the population.16
Somehow the different conditions altered the growth of all the individuals
exposed to them, and if such an effect were inherited it would become
cumulative. If the external conditions were seen as the chief factor in di-
rected variation, closely related species might be expected to react to the
same conditions in the same way. But one could hardly expect widely
different species to follow the same trend, and even within a single species
12. Ibid., pp. 134-139.

13. Sec the discussion of Cope's work in Kellogg, Darwinismtoday(n. 1), pp. 28 5288; and Richard
Swann Lull, Organic evolution,rev.ed. (New York, 1929), chapter 12.
14. Henry Walter Bates, 'Contributions to an insect fauna of the Amazon valley. Lepidopteia:
Heliconidae," Trans. Linn. sec. Land., 1862, 23, 495-566, p. 514.
15. See Ludwig Plate, Uber Bedeutung its Darwin'schen Selectionsprinzip und Problemc icr Artbildung
(Leipzig, 1903), p. 187. Julian Huxley, Evolution (n. 3), p. 500 notes the importance of this concept
and attributes the name orthoselection to Plate, but lists only a later work.
16. Darwin, Origin of species (n. 11), pp. 132-134.
44 Journal of the History of Medicine : January 1979
orthogenesis would only continue in the same direction as long as the en-
vironment changed consistently. Eimer himself maintained that external
factors were the stimulus that produced orthogenesis, and insisted that
since orthogenetic changes would not necessarily be adaptive they should
be considered quite separate from the adaptive trends of Lamarckism. Al-
pheus Packard, the American entomologist and prominent neo-Lamarck-
ian, disagreed and showed that Lamarck himself had postulated such a
direct effect of the external conditions in addition to use and disuse.17 But
Packard's criticism was really a quibble over nomenclature, because even
if Lamarck accepted both mechanisms, there was still a need to distinguish
between them. Most neo-Lamarckians concentrated on the adaptive effects
of use and disuse, so Eimer's insistence that orthogenesis be treated as a
separate effect probably served a valuable function.
Even in cases where the environment was the stimulus producing the
variation, it had to be considered as acting upon the existing constitution
of the organism. The constitution might predispose the organism to react
to stimuli in certain ways, and such a predisposition might be the main
directing factor. Darwin was aware of cases in which variation appeared
to be directed from within, as when a number of forms varied in the same
direction, even when not exposed to the same conditions.18 Thomas Henry
Huxley also thought that there might be constitutional limits on the direc-
tion of variation.19 In such cases the environment might still act as the
stimulus to elicit change, but the pattern of development would be prede-
termined by the internal constitution of the species. Whatever the stimulus,
the same changes would be produced in all species with the same predis-
position, so that a whole group might undergo a parallel and consistent
orthogenesis. Thus the scope of the trends to some extent determined the
mechanism that seemed most suitable for explaining them. The particular
reactions of individual species to changes in the local conditions suggested
that the environment was the chief directing factor, while large scale trends
pushing whole groups of species consistently in the same direction over
long periods implied some kind of internal direction. To some extent the
story of Eimer and his followers revolves around the tension between these
two possible explanations of orthogenesis. Both interpretations, however,
17. Alpheus Packard, Lamarck, thefounder of evolution: his life and work. With translations of his writings
on organic evolution (New York, 1901), pp. 408-409.
18. Darwin, Origin of species (n. n ) , p. 159.
19. Thomas Henry Huxky, 'Evolution in biology1 (1878), reprinted in Collected Essays, n, Dar-
winiana (London, 1893), 187326, p. 023.
could be pushed to an extreme where they eliminate the need for selection
by making variation itself the driving force of evolution.
Eimer championed external factors as the chief influence on orthogen-
esis. Yet he also made sweeping claims about the extent and unity of or-
thogenetic trends that imply a strong degree of internal direction. His
followers picked up both possible interpretations. A number of attempts
were made to demonstrate the direct action of local conditions on a species.
But as time went on, the internal factor became generally more popular.
The wider the scope of the trends postulated, the more difficult it was to
believe that external causes could provide enough consistency to account
for the phenomenon. This was true of Eimer's later work and more espe-
cially when the paleontologists began to describe trends that continued
over vast periods of time. In addition, Weismann's theory of the germ
plasm and the rediscovery of Mendelism both tended to eliminate the pos-
sibility that external effects on individual growth could be inherited. In
the end the most plausible mechanism that could be suggested for ortho-
genesis was some property of the germinal material itself that could con-
trol variation. Weismann proposed his theory of germinal selection to
allow for orthogenesis, although there were other approaches that could
also be explored.
One important argument for external causes was that they could be seen
as a purely mechanistic factor carrying no implication that the trends were
goal directed. Darwin had insisted on the random nature of most variation
because he wanted to eliminate teleology. When the American botanist
Asa Gray tried to retain a role for divine benevolence by suggesting that
variation was supernaturally directed toward useful ends, Darwin pro-
tested that such divine control would eliminate all hope of a scientific ex-
planation.20 The majority of naturalists eventually came to accept Darwin's
view, and even neo-Lamarckism became only an alternative naturalistic
explanation of adaptation.21 But what about nonadaptive trends? The
British physiologist William Benjamin Carpenter argued that nonadaptive
patterns, for instance in the evolution of the foraminifera, were evidence
20. For Darwin's public critique of Gray's position see the conclusion of his Variations of animals and
plants under domestication, and ed. rev., 2 vok (London, 1882), n, 427-408. Gray's articles on theology
and evolution are collected in hij Darwiniana: essays and reviews pertaining to Darwinism (1876, reprint
ed., Cambridge, Mia., 1963).
21. Edward J. PfeuTer has suggested that American neo-Lamarckism was associated with natural
theology; see his "The genesis of American neo-Lamarckism,' his, 1065, $6,156-167, but I have argued
against this elsewhere, see my 'Edward Drinker Cope and the rhanging jtructurc of evolutionary
theory,' lsis, 1977, 6t, 249-265.
\6 Journal of the History of Medicine : January igjg
of supernatural design aimed at the production of complexity and beauty.22
From an entirely different background, Karl von Nageli used trie presence
of nonadaptive structures to discredit natural selection and substitute for it
an inner perfecting force or Vervollkommnungskraft.23 Eimer stressed ex-
ternal factors in orthogenesis precisely because he wanted to dissociate
himself from Nageli's use of such an apparently mysterious inner force.
Most later support for internal factors came from workers who insisted
that there must be some natural mechanism involved to eliminate design.
But the proponents of orthogenesis were always on the defensive against
teleology, as though they were aware of the constant temptation to read a
meaningful goal into their trends.
BIMER'S EXPOSITION OF ORTHOGENESIS
In 1869 Eimer completed his studies under Kolliker, who was known as a
supporter of directed variation and sudden mutation in evolution.24 Al-
though Eimer seldom referred to Kolliker in his later writings, his first
important publication in 1874a study of the color variation of the rock
or wall lizards of Capri, particularly the distinctive blue-black variety of
the Faraglioni rocksdealt with a case of possible directed variation. These
unusual lizards later attracted the attention of a number of naturalists, in-
cluding Alfred Russel Wallace, but Eimer's study was one of the most
thorough.25 In 1874 his attitude toward natural selection was ambivalent.
His observations of the Capri lizards seemed to indicate that a new variety
or incipient species could appear in a particular locality, and according to
Eimer they also suggested that the new form was produced suddenly by
22. William Benjamin Carpenter, "The argument from design in the organic world,' reprinted in
Nature and man: essays scientific and philosophical (New York, 1889), pp. 445-463. On the role of this
idealist concept of design see my 'Darwinism and the argument from design: suggestions for a re-
evaluation,' J. Hist. Biol,, 1977, 10, 29-43.
23. Nageli s theory was proposed in his Entstehung del Bcgriffes der noturhistorischen Art of 1865 and
then developed in his Mechanische-physiologische Theorie des Abstommungslehre (Munich and Leipzig,
1884). A summary of the latter was published as A mechanical-physiological theory of organic evolution
(Chicago, 1898). Nageli's objections to selection are quoted in Kellogg, Darwinism today (n. 1), pp.
62-64.
24. The most accessible account of KSlliker's Vber die Darwin'sche Schopjungstheorie of 1864 is T. H.
Huxley's 'Criticisms of the "Origin of species," ' (1864) in Huxky, Collected essays, n (n. 19), pp.
80-106.
25. Theodor Eimer, Zoologische Studien auf Capri, n. Lacerta muralis caerula, ein Batrag zur Dar-
win'schen Lehre (Leipzig, 1874). A brief note of this work appeared in English: 'Lacerta muralis cae-
rula: a contribution to the Darwinian theory,' Ann. & Mag. Nat. Hist., 1875, 16, 234-235. See also
Eimer, Organic evolution (n. 8), pp. 20-21. For Wallace's comment see his 'Remarkable local color-
variation in lizards,' Nature, 1878-79, ig, 4. See also A. Leith-Adam's note under the same title, ibid.,
P- 53-
internal causes without reference to utility. But at this early stage in his
career, Eimer was prepared to admit that only useful variations would be
able to establish themselves permanently in the species, and he was actually
criticized for stating that the color of the Capri lizards had adaptive value.26
Directed variation was thus an addition to the selection theory rather than
an alternative to it.
In the 1880s Eimer began to adopt a more hostile attitude toward selec-
tionism. During this decade Weismann gradually elaborated his germ
plasm theory and began to argue that selection was the only effective evo-
lutionary mechanism. The original form of Darwinism had never been so
exclusiveeven Darwin himself accepted use inheritance and directed
variation. Weismann's neo-Darwinism polarized opinions by creating an
'all or nothing' attitude toward natural selection. Having decided from the
first that selection was not the whole story, Eimer seems increasingly to
have thought that he now had to oppose selection in order to retain a role
for alternative mechanisms. To demonstrate that there were large areas of
evolution which could not have been caused by selection, Eimer began to
take up Nageli's argument that all species possess characters of no adaptive
value. These would have to be explained by directed variationthe mech-
anism later to be called orthogenesis. Eimer also began to insist that even
the adaptive characters can be explained more readily by Lamarckism than
by natural selection. These objections to neo-Darwinism were finally col-
lected together in Die Entstehung der Alien of 18 8 8, the work that was trans-
lated into English as Organic evolution. This was a compendium of material
drawn from many sources, supplemented by Eimer's own observations
on various kinds of animals. The Capri lizards in 1888 were used as evi-
dence against selectionism and in favor of directed variation as the driv-
ing force of evolution. But by 1888 Eimer had begun the study of the
colors and patterns on the wings of butterflies which was to become his
main line of evidence for orthogenesis. In 1889 he published a preliminary
report on this work 27 and used the same material as the basis of an address
in 1895. The address was then expanded to form the first chapter of Eimer's
26. Eimer, Lacerta muralis caaula (n. 25), p. 36. See Henry Hillyer Gigioli, 'Colour variation in
lizardsConican herpetology,' Nature, 1878-79,19, 97; A. Emit, 'Local colour-variation in lizardj,'
Nature, 1879, 20, 290-291; and J. von Bedriaga'i note under the ume title, Nature, pp. 480-481. Von
Bedriaga refers to hii Ueber die Entstehung der Farben bei den Eidtchsen of 1874 and Die Faraglioni-
Eidechse of 1876. He argued that the Faraglioni rocks were of a light color, hence the blue-black of the
lizards could not be camouflage as Eimer had claimed. Nor was there any indication that the color
was a protection against the sun on the exposed rocks, since it was not shown by similar lizards
inhabiting deserts.
27. Theodor Eimer, Artbildung und Verwandschaft bei dm Sduneturlingen (Jena, 1889).
major work on orthogenesis, that in 1897 formed the second volume of
his Die Entstehung der Arten.
In the later works devoted purely to orthogenesis, Eimer developed
even further the claim that all species have nonadaptive charactersindeed
he now seemed to be making a positive effort to play down die significance
of adaptation in evolution. The great fault of Weismann's neo-Darwinism
(Eimer called it 'pseudo-Darwinism') was that it had to treat nature as a
completely utilitarian system in which every character had a definite adap-
tive value. Following earlier critics of Darwin such as St. George Jackson
Mivart, Eimer pointed out that even those characters that do acquire some
adaptive value must have passed dirough incipient stages when they would
not be useful.28 In Eimer's own field of animal coloration, he thought the
utilitarian element had been greatly exaggerated. It was always possible
for the neo-Darwinian to make a wild guess about die adaptive value of a
particular color-scheme, but such guesses could never be substantiated and
were seldom very plausible to the experienced observer. After exerting all
his ingenuity to show how the colors on, say, a snail's shell might serve as
camouflage, the Darwinian was still unable to account for die colors on
the inside of die shell.29 All such cases could be explained by assuming diat
variation could be directed by forces diat might be stimulated by the en-
vironment, but which were in no way an adaptive response to it. In the
end Eimer was so convinced of the prevalence of directed variation that he
claimed there was no random variation at all.30 All changes were rigidly
directed by die laws of ordiogenesis, or by Lamarckism. Eimer's observa-
tions carried the heavy burden of his effort to prove such extreme claims.
The observations on butterflies fell into two categories. By synmesi/ing
die reports of odier workers, Eimer was able to establish a plausible claim
that temperature had a direct effect on coloration. Max Standfuss had
found diat when pupae of Papilio machaon were exposed to artificial heat
in die laboratory, die resulting butterflies resembled diose normally found
in die soudiern part of die species' range.31 Butterflies that appeared during
28. Eimcr, On orthogenesis (n. a), p. 4. See S t George Jicloon Mivart, O B the genesis ofspedts (Lon-
don, 1871), chapter 2.
29. Eimer, Orthogenesis (n. a), p. 10.
30. Ibid., pp. 24-35. See alio Theodor Eimer, Die Entstehung der Arten . . . xweiter Theil. Ortho-
genesis der Schmetterlinge: ein Beweis bestimmt gerithteter Entwickclung und Ohnmacht der naturlichen Zutht-
wahl bei der Artbildung (Leipzig, 1897), p. 16.
31. Ibid., p. 36; Orthogenesis der Schmetterlinge, pp. 26-27. The latter work alto hai a whole chapter
on environmental influence!, pp. 388-456. Standfuu' work wai fummarized in English: 'Synopsil of
experiment! in hybridization and temperature tnarfr with Lepidoptera up to the end of 1898,' Ento-
mologist, 1000,33, 161-167,183-293. 340-348 and IOOI, 34, 11-13, 75-84.
Bowler : Theodor Elmer and Orthogenesis 49
the hottest part of the summer tended to have the same southerly character.
In such cases the external conditions produced a direct modification of in-
dividual growth. Although no studies were made to confirm that such
variations would be inherited, their inheritance was widely assumed to
occureven Weismann accepted seasonal dimorphism in butterflies as a
genuine instance of directed variation.
Eimer's own observations were meant to establish the existence of far
more extensive orthogenetic trends, but they used a different approach.
Eimer arranged the species within the genus Papilio to give an apparently
regular sequence of color and pattern modifications, a sequence which was
supposed to be a record of the group's orthogenetic evolution. The basic
sequence was from longitudinal stripes to spots, cross-stripes and finally
uniform color, with the changes starting at the back of the wing and
spreading forward.32 Several species had eleven longitudinal stripes: Papilio
alebion, P. paphus and P. glycerion. Of these Eimer wrote: 'Now these lon-
gitudinally striped butterflies give, as I am becoming more and more con-
vinced, the fundamental form of marking in all diurnal butterflies.'33 The
number of stripes became reduced successively in Papilio eurymedon, P.
tumus, P. alexanor and P. machaon down to a more or less uniform color in
P. asterias. Spots appeared in P. machaon and became increasingly promi-
nent in P. xuthus and P. xulanthus. When two species exhibited roughly
similar patterns, Eimer believed that the similarity resulted from two sepa-
rate lines of evolution that had reached the same point on the sequence of
orthogenetic development. This phenomenon of parallel development ex-
plained the resemblances that the Darwinians attributed to mimicry.34 Ei-
mer even seems to have thought that varieties within the same species owed
their relationship to parallel development rather than common descent.
To Eimer it was obvious that the sequence color and pattern of modern
species of butterflies must represent the past evolution of the group:
On the plates of my butterflies the formation of species and the laws of evolution
can be read direcdy from the wings. The markings and colorings of the same
are so many letters speaking a clear and forcible language that no one who
wants to know the truth can misunderstand. Like the leaves of an open book
the written characters on the wings of our butterflies show their past and present
history.35
32. The bws of orthogenesis ire summed up in Pimw, On orthogenesis (n. 2), p. 26 and Eimer,
Orthogenesis der SchmetteHinge (n. 30), pp. 18-20.
33. OH. (n. 2), pp. 37-38; (n. 30), p. 28.
34. Ibid. (n. 2), p. J2; (n. 30), p. 38.
33. Oid. (n. 2), p. 37; (n. 30), p. 28.
But to anyone familiar with the principles of Darwinism, Eimer's inter-
pretation must have seemed quite arbitrary. Eimer could never explain
why he was convinced that the longitudinal stripes represented the starting
point, nor why the others must have followed in so regular a sequence.
The Darwinian would ask how we can be sure that any modern form still
retains the characters from which die group began; each species may have
changed in its own way and there is no reason why a pattern made by ar-
ranging the modern forms in what looked like a regular order must cor-
respond to the past evolution of die group. The only sure guide to the
exact nature of earlier forms was paleontology. Eimer was aware of the
fossil evidence for ordiogenesis, citing in particular Alpheus Hyatt's work
on the parallel and regular developments of the cephalopods.36 But pale-
ontology was useless as a guide to the patterns of butterflies, and Eimer
was convinced diat his own arrangements of the living species were more
valuable than fossils as guides to die evolutionary process. He remained
completely unaware of the arbitrary nature of his assumptions. It is hardly
surprising that Weismann refused to take this side of Eimer's work se-
riously.
Whatever the value of Eimer's observations, he was able to work out
the mechanism of orthogenesis in some detail. Superficially, the theory
was presented as an equal combination of the two factors: internal and
external. 'The causes of definitely directed evolution are contained, accord-
ing to my view, in the effects produced by outward circumstances and
influences such as climate and nutrition upon the constitution of a given
organism.'37 The constitution of die organism dictated the 'laws of growth'
which would respond in a specific way to each kind of external stimulus.
Divergence of one species into many would be possible because exposure
to different conditions in different geographical areas would change die
constitution of die original species in various ways, and each new form
would respond uniquely to future stimuli. Eimer wrote:
In my view development can take place in only a few directions because the
36. See for instance ibid. (n. a), p. 3 and p. 12; (n. 30), p. 5 and p. 19. Eimer cites Hyatt'i Genesis of
the Arictidat (Washington, 1889). Eimer'i lawj of orthogenesu included Hyatt and Cope'i law of the
acceleration of growth: when a new character appears, earlier characters must be compressed back
into embryological growth to make room for it; tee Bowler (n. 21) and Stephen Gould, Ontogeny
and phytogeny (Cambridge, Mas)., 1977), pp. 85-96. Hyatt and Cope both came to accept that use-
inheritance directed the addition of new stages in progressive evolution. But Hyatt also traced out
sequences of degeneration in the cephalopods which he claimed were the result of exposure to advene
conditions, and these later became known as classic cases of orthogenesii in paleontology.
37. Eimer, Orthogenesis (n. 2), p. 22; Eimer, Orthogenesis der Schmetterlingc (n. 30), p. 13. See also
the passage translated in Kellogg, Darwinism today (n. 1), p. 285.
Bowler : Theodor Eitner and Orthogenesis 51
constitution, the material composition of the body, necessarily determines such
directions and prevents indiscriminate modification.
But through the agency of outward influences the constitution must gradually
get changed. The organisms will thus acquire more and more physiological in-
dividuality and respond to outward influences more and more in a manner har-
monising with their specific individualityand so new directions of develop-
ment will be produced.38
On the other hand, if changed conditions affect the whole of a species, a
new form may appear in the midst of the original population. The species
will divide itself by 'genepistasis'the tendency of the more sensitive in-
dividuals to progress to the next stage of variation ahead of their fellows.
According to Eimer this again illustrated the superiority of orthogenesis
over Darwinism, since it explained the appearance of new forms without
isolation.39
Eimer's ideas may appear strange to the modern reader, but they were
consistent and even plausible, given the basic assumption that such varia-
tions were hereditable. The theory was also presented as completely mech-
anistic: although there was a directing force from within the species this
was provided by the physiological laws of growth, not by a mysterious
'perfecting principle' such as Nageli had advocated. The fact that ortho-
genesis sometimes produced degeneration showed that the trends were not
directed toward any meaningful goal and did not require a teleological
explanation.40 Eimer also insisted that his inclusion of external factors
marked a crucial distinction between his own mechanistic approach and
Nageli's vitalism.41 Standfuss' experiments clearly showed that the nature
of the external conditions played a fundamental role in determining how
the laws of growth were modified. If the laws of growth alone directed
the whole course of evolution, the history of life would be completely
predetermined, and to accept this one would have to endow life with more
or less supernatural powers. Only by insisting that orthogenesis was pro-
duced by the environment's interference with the previously established
laws of development could it be made a scientifically respectable hypoth-
esis. The balance between the internal and external factors was thus a fun-
damental element of the image that Eimer's theory presented to the world.
38. Eimer, Orthogenesis (a. a), p. aa; Eimer, Orthogenesis da Schmetlerlinge (n. 30), p. 13.
39. Ibid. (n. a), p. 30; (n. 30), p. at. Note that Eimer also saw tuch changes 35 saltations, although
he believed that the sudden chaagei were relatively tmall ones.
40. Ibid. (n. a), p. 40; (n. 30), p. 29.
41. Ibid. (n. a), p. isn and pp. 20-23; (n. 30), pp. 8-on and pp. 13-16.
Unfortunately Eimer was unable to maintain this relatively sophisticated
position throughout his writings. Although in theory the environment
played as important a role as the laws of growth in determining the direc-
tion of orthogenesis, we often find Eimer implying that the internal factor
exercises far more control. His own interpretation of the butterflies indi-
cated that there were only a few basic directions of variation open to the
group, whatever the conditions to which the individual species might be
exposed. At one point he admitted that the development he postulated
seemed to have occurred under conditions opposite to what would have
been expected from the experiments on temperature.42 In effect, the pat-
terns of variation were somehow predetermined by the nature of the or-
ganisms, and the environment could do little to produce significant diver-
gence. The evolution of the butterflies was along a number of parallel lines
moving at different speeds through the same pattern of development.
Eimer was even prepared to argue that the same pattern of color variation
was predetermined not only for the butterflies but also for the whole of
the animal kingdom. It could be observed in other insects, in molluscs and
even (as with the lizards) among the vertebrates.43 The same rigid limits
on variation must hold for every other character in both the animal and
the vegetable kingdoms:
Orthogenesis is a universal law. It holds, as I have long insisted, not only for the
markings but also for the other morphological characters of animals, as also for
those of plants. Even in the latter, as my personal observation has convinced me,
the markings of the blossoms and the shape of the leaves follow, in rigorous
conformity to law, a few definite directions.44
The 'law' of orthogenesis had obviously become for Eimer something
more than a balance between the physical constitution of the organism
and the external stimuli. The whole evolution of life had been able to
unfold only along certain paths, with true divergence only being possible
within the combinations allowed by the trends. Elsewhere in Eimer's writ-
ings there are exaggerated eulogies of the principle of the unity of nature
43. Ibid. (n. a), p. 33; (n. 30), p. 38.

43. Ibid. (n. a), pp. 7-9; (n. 30), pp. 3-4.
44. Ibid. (n. a), p. 31; (n. 30), p. 14. Compare thii quotation with that given earlier (a. 38), where
Eimer admit] that exposure to different conditions will generate new Una of developmentthe two
passages are only a few pages apart, yet have a completely different emphasij. Note, however, that
even when discussing evolutionary divergence, Eimer insisted that it exhibited a degree of regularity
and simplicity that could not be explained as the result of random influences, see ibid. (n. a), pp. 34-
2j; (n. 30), p. 16.
and even a favorable reference to the philosophy of Lorenz Oken.45 For
all his efforts to present orthogenesis as a fashionably mechanistic theory,
Eimer had not in fact been able to break away from the old spirit of
Naturphilosophie. Nature may be a system of material interactions, but she
is also a harmonious whole, programmed to unfold in a fixed pattern.
Admittedly the trends are not supposed to run toward a meaningful goal,
and strictly speaking Eimer had eliminated teleology, but the system de-
manded a unity of overall structure that was incompatible with most post-
Darwinian world views. Small wonder that Weismann and many other
naturalists were suspicious of Eimer's more general claims.
WEISMANN'S CRITIQUE OF ORTHOGENESIS
In 1888 Eimer proclaimed his opposition to Weismann's neo-Darwinism

and reinforced his opposition to Weismann in his later writings on or-
thogenesis. Weismann seldom wasted much effort in replying to Eimer
directly, thereby fuelling the latter's belief that he was being systemati-
cally snubbed. But there is evidence that Weismann took the question
of directed variation very seriously indeed, and was constantly forced to
modify his position in order to accommodate it. What he was not pre-
pared to do was tolerate Eimer's final vision of an evolutionary process
dominated by a few broad orthogenetic trends. Directed nonadaptive vari-
ation might be accepted as long as it was treated as a minor addition to
natural selectionmuch as in Eimer's first discussion of the Capri lizards.
This meant that the orthogenetic trends must be limited to particular de-
velopments by individual species, with most variations still being random
and subject to selection. The more limited and plausible of Eimer's ex-
amples could be accepted as genuine exceptions to the rule of random
variation, while his overall world view was ignored as a throwback to the
old idealist philosophy of nature.
The first of Weismann's works to gain wide recognition was, in fact, a
collection of studies devoted to directed variation: his Studien zur Descen-
denztheorie of 1875. In the preface to the English translation of this work,
Webmann explained that many German-speaking biologbts had rejected
selection in favor of directed variation, citing Nageli, Kolliker, and their
followers as examples.46 Weismann's own purpose was not to eliminate
4J. Eimer, Organic evolution (n. 8), pp. 409-410 and p. 433; Elmer, Entstchung iet Arten, pp. 440-
443 and p. 460.
46. Auguit Weinnann, Studies in the theory of descent, trans. Raphael Meldola, 2 volj. (188a; reprint
ed.. New York, 1975), 1, xvii-xviii.
the possibility of directed variation but to show that it occurred only in a
limited form that could not displace selection as the main evolutionary
mechanism. In the course of his observations he actually referred to Eimer's
study of the Capri lizards as another example of the limited kind of di-
rected variation that he was himself finding among insects.47
In a study of seasonal dimorphism among butterflies, Weismann was
forced to admit that those individuals born early in the season differed
systematically from those that appeared later. As in Standfuss' experiments
the difference could be reproduced by exposure to artificial heat, and hence
must be due to the direct action of the external conditions. Weismann still
believed that changing conditions normally stimulated only random vari-
ation, but there was no doubt that in a few cases the whole species might
react in the same way. '. . . In butterflies at least, all the individuals of a
species respond to the same external influences by similar changes, and . . .
consequently, the changes brought about by climatic influences take a
fixed direction, determined by the physical constitution of the species.'48
Weismann assumed that such changes could be inherited, and hence that
if exposed to new conditions for long enough the species would be per-
manently changed. A simple reversal of the conditions would not recreate
the old form, since the new species would have its own unique reaction to
any stimulus.49
Seasonal dimorphism involved different changes in each species, but
there were other cases in which a more general trend seemed to affect a
number of closely related forms. In some caterpillars there was a tendency
for spots produced on one segment to spread by repetition along the whole
animal. The original spot might have served some purpose to frighten
enemies, but the extra ones had no such value and so could not have been
produced by selection. Such a tendency was particularly strong in the
genus Smerinthus, of which Weismann wrote:
How is it then that three species vary independently of each other in an analo-
gous manner? I know of no other answer to this question than that similar
variations must necessarily arise from similar physical constitutionsor, other-
wise expressed, the three species have inherited from an unknown parent species,
devoid of spots, not dais last character itself, but a physical constitution, having
a tendency to the formation of red spots on die skin.50
47. Ibid., I, 361.
48. Ibid., I, 108.
49. Ibid., 1, 104.
50. Ibid., 1, 360. O n Weismann's efforts to explain the coloration o f caterpillars lee Gould, Ontogeny
(n. 36), pp. 103-109.
At this point Weismann cited Eimer's observations on the wall lizard as
providing evidence of an analogous phenomenon. Here the principal cause
of directed variation appeared to be the internal predisposition acting in-
dependently of the external conditions.
In later years Eimer liked to point out that Weismann's original views
had been in agreement with his own. The above quotations show that
there was some truth in Eimer's claim, but in fact Weismann always tried
to minimize the significance of the few apparently incontrovertible cases
of directed variation. Whereas Eimer went on to claim that all evolution
was directed according to a few fixed trends, Weismann's main concern
was always to eliminate the possibility of any significant control of evolu-
tion by an internal force. He argued that even where parallel variation
occurred, there was not enough regularity in the phenomenon to suggest
large-scale internal predisposition.51 Variation was normally random and
selection the main driving force of evolution. Only rarely was a whole
speciesand even more rarely a group of related speciesforced to vary
consistently in the same direction.
In the beginning Weismann had accepted both external and internal
factors governing variation, but as he began to develop his celebrated
theory of the germ plasm in the course of the 1880s Weismann had to
rethink his position.52 The germ plasm was supposed to be independent of
the somatic cells of the body, hence no effect of the external conditions on
the growth of the individual (adaptive or nonadaptive) could be inherited.
In 1886 Weismann argued that individual differences within the species
could not have been produced directly by the environment except during
the earliest phase of evolution before the development of sexual reproduc-
tion.53 In 1892 he clarified this statement and tried to return to his original
position. In a sense all the differences among individuals did date back to
the earliest period of evolution, but in the course of time each individual
germ plasm has responded differently to changing conditions and new
factors have constantly been introduced into evolution. External con-
ditions could thus affect the germ plasm directly, not via the soma.Much of
the effect of external conditions resulted in merely random variation, but
occasionally many individuals within the species might possess a germinal
ji. Webmann, Theory of descent (n. 46), 1, 363.

50. On the development of the germ plasm theory, see for imtanrr Frederick B. Churchill, 'August
Weismann and a break from tradition,' J. Hist. Bio/., 1968, 1, 91-112.
53. August Weiimann, The significance of sexual reproduction in the theory of natural selection'
(1886) reprinted in hij Essays upon heredity and kindred biological problems, ed. Edward B. Poulton et
al., 2 vols. (Oxford, 1891-92), I, 284-286.
constitution that reacted in the same way to a particular external change.54
Thus directed variation would again be the product of a combination of
external factors acting on the internal (germinal, not physiological) con-
stitution of the organism.
Soon, however, Weismann seems to have realized that in allowing the
direct action of the environment upon the germ he was dangerously com-
promising his basic idea of the independence of the germinal material. If
directed variation occurred, it would have to be attributed to some factor
within the germ itself. Yet Weismann's basic concept of the germ plasm
as composed of a collection of 'determinants' for each character did not
encourage the belief that die germ plasm could be structurally predisposed
to vary more easily in certain directions. Instead Weismann proposed in
1895 a n active process within the germ that was potentially capable of
developing any determinant as a source of directed variation: germinal
selection. Having for years brushed off the objections to natural selection
(which he now called 'personal selection'), Weismann's faith in his new
germinal mechanism was strong enough for him at last to admit that the
traditional Darwinian theory was inadequate.55 His new addition to the
theory followed the lead of Wilhelm Roux, who recognized that a state of
struggle could exist among parts of the body.56 Germinal selection trans-
ferred this internal struggle to the germ plasm itself: the determinants
could compete with one another and hence be subject to selection. Only
so much nourishment was available for the germ plasm and all the deter-
minants must compete for their share. Personal selection picked out not
only those individuals with the appropriately developed character, but also
those whose germ plasm contained a strong determinant for producing
that character. Such a strong determinant would then predominate in the
struggle for nourishment and its success would result in further variation
in the same direction.
For as soon as personal selection favors the more powerful variations of a deter-
minant, the moment that these come to predominate in the germ plasm of the
species, at once the tendency must arise for them to vary still more strongly in
the plus direction, not solely because the zero-point [of variation] has been
pushed further upwards, but because they themselves now oppose a relatively
more powerful front to their neighbors, that is, actively absorb more nutriment,
54. August WaiTnann. The germ plasm: a theory of heredity, tram. W. Newton Parker nd Harriet
Ronnfeldt (London, 1893), pp. 401-403.
55. Weumann, 'On germinal selection' (n. io), pp. 263-264.
56. Wilbelm Roux, Der Ksmpjier Theile im Organismus (Leipzig, 1881).
and upon the whole increase in vigor and produce more robust descendants.
From the relative vigor or dynamic status of the germ-plasm, thus, will issue
spontaneously an ascending line of variation, precisely as the facts of evolution
require.57
In the opposite case of an existing character that was rendered useless be-
cause of changing demands on the species, the relaxation of personal selec-
tion would allow the emergence of weak determinants which would then
be gradually eliminated through germinal selection.58
In his original account of germinal selection, Weismann wrote as though
utility determined the initial direction of each line of variation. But the
determining role of utility still did not eliminate the anti-Darwinian argu-
ments based on the existence of nonadaptive characters. To deal with these
Weismann in 1902 gave a somewhat different account of germinal selec-
tion and retreated even further from his original position. He now ad-
mitted that selection could not explain the incipient stages of organs that
became useful only when well developed, and that even some quite prom-
inent characters had no utilitarian value.59 It must thus be possible for any
determinant to gain an advantage by chance in the struggle for nourish-
ment, thereby establishing a line of directed variation with no real value
to the species. Weismann still insisted that such a process rarely achieved
significant proportions and that it could never produce positively harmful
characters. However, he was forced to admit that in some cases the results
may have been disastrous for the species, as with the Irish elk, which was
widely supposed to have been driven to extinction by the over-develop-
ment of its antlers.60 Here germinal selection may have produced a useless
character which then became harmful because of some sudden change in
the external conditions. However rare Weismann supposed such instances
57. Weismann (n. 10), pp. 275-276.

58. Ibid., pp. 273-274. Weismann regarded the reduction of iw*li* organs as even more crucial
than the positive case. Originally he had explained such reductions solely in terms of the relaxation of
selection (panmixia), which allowed the weaker individuals to proliferate and hence reduce the aver-
age size of the organ. This did not seem to explain the almost complete elimination which has oc-
curred in some cases, and Weismann himself was now admitting the inadequacy of his earlier views.
59. August Weismann. The evolution theory, trans. J. Arthur Thomson and M. R. Thomson, 2 vok
(London, 1904), n, 132-13+. The German edition appeared first in 1002.
60. Ibid., p. 3 j8. The 'overdevelopment' theory of extinction became quite popular among twen-
tieth century paleontologist supporters of orthogenesis such as Henry Fairfield Osbom. Weismann
cited Ludwig Doderkin as its originator, see Ludwig Doderlein, 'Phylogenetische Betrachtungen,'
Biologisches Centralblatt, 1888, 7, 394.-402. On this particular case of supposed overdevelopment see
Stephen Jay Gould, The origin and function of "bizarre" structures: antler size and skull ti1** in the
"Irish elk," Megaloccros gigantau,' Evolution, 1974, 28, 191-220.
to be, it is clear that he no longer saw utility as the sole driving force of
evolution.
Weismann now hailed germinal selection as a compromise between the
systems of Nageli and Darwin61a claim which reveals how well aware
he was of the extent to which he had been forced to compromise with the
idea of an internal directing force. And (although Weismann never ad-
mitted it) he had also made a partial compromise with Eimer's views.
Both had begun by attributing orthogenesis to the inherited effects of the
external conditions, but had then gone on to emphasize the role of internal
direction. Yet there was still a world of difference between them. For
Eimer, the laws of growth dictating the patterns of orthogenesis were fun-
damental unifying principles linking all evolution. But Weismann con-
ceived the directing power of the germ in a way that could provide no
such unity even within related forms. Germinal selection could produce
directed variation, but each line of development would be a unique process
going in its own direction, started by chance but ultimately limited by
utility. There was no structure within the germ which could impose a pre-
determined limit on variation, since each determinant functioned indepen-
dently of all the rest. Perhaps Weismann might have explained parallel
variation in related species by supposing that some determinants could be-
come dominant more readily than others, but he never took this course
and now ignored cases such as the Smerinthus caterpillars. No doubt Eimer
exaggerated the extent to which the unfolding of orthogenesis could be
predetermined, but Weismann had gone to the opposite extreme of com-
pletely eliminating the regularity of development that to many seemed
the best evidence for directed variation.
BRITISH AND AMERICAN REACTIONS
In 1897 Eimer published a vigorous critique of germinal selection.62 In his

opinion the mechanism still did not concede a sufficient role to directed
variation and merely served to indicate the vulnerability of Weismann's
original position. The whole germ plasm theory was in any case an unjus-
tified speculation in the eyes of any supporter of Lamarckism and exter-
nally stimulated orthogenesis. Germinal selection was merely aflimsysu-
perstructure erected on the foundation of what was already an extravagant
hypothesis. But Eimer died in the following year, leaving it for others to
press home the claims of orthogenesis as an evolutionary mechanism. The
61. Weinnann, Evolution theory (n. 59), 1, 33X
62. Eimer, Enwehung da Arttn (n. 30), pp. 50-97.
Bowler : Theodor Einter and Orthogenesis 59
widely differing interpretations of directed variation offered by Weismann
and Eimer formed important components of the widespread debate over
the validity of natural selection that broke out around the turn of the cen-
tury. Eimer's own work was taken seriously by several British and Amer-
ican naturalists, whose reactions illustrate the unresolved conflict in Eimer's
writings between internal and external control of variation. In either case,
though, such reactions illustrate the opposition that had arisen in some
quarters to the utilitarian commitment of neo-Darwinismopposition to
which Weismann himself made concessions with germinal selection.
Eimer's claim that directed variation was only produced in response to
environmental stimulus was calculated to appeal to any member of the
neo-Lamarckian school. If they could not follow Eimer's efforts to limit
the role of adaptation in evolution, most neo-Lamarckians still agreed that
use-inheritance would have to be supplemented to at least some extent by
a mechanism for producing nonadaptive variation. Thus Edward Drinker
Cope's classic summary of the Lamarckdan positionThe primary factors of
organic evolution of 1896included short but by no means insignificant sec-
tions on directed variation. Cope was not averse to the search for regulari-
ties in evolution; as one of the original proponents of the 'law of accelera-
tion of growth' he had always maintained that evolution proceeded by the
regular addition of stages to individual development.63 He had gradually
come to admit that use-inheritance, guided by the existing habits of the
species, was the factor which most often determined what new stages were
to be added. But Cope always believed that there were at least some char-
acters with no adaptive value, and in his studies of color variation among
North American reptiles he obtained results that he compared directly
with those of Eimer on the wall lizard.64 Cope certainly held that such
variation exhibited regularities, in the sense that the varieties within a spe-
cies could be arranged into an orderly sequence.65 But he assumed that for
the most part these sequences corresponded to the geographical distribu-
tion of the varieties, with the regularity of variation being produced by
the gradual change of factors such as temperature and humidity over a
63. On the development of Cope's views jee Bowler, 'Edward Drinker Cope' (n. 31) and Gould,
Ontogeny (n. 36). Originally Cope had itrongly opposed the utilitarian view of nature, but he soon
abandoned thii poiition as he came to accept the significance of use-inheritance.
64. Edward Drinker Cope, The primary factors of organic evolution (Chicago, 1896), pp. 13-45. Cope
cites a number of his own studies on the milt mitr, Osceola ioliata and various species of lizards. See
also Packard, Lamarck (a. 17).
65. Ibid., p. 39. See for instance the diagram representing the varieties of the milk snake.
wide geographical area.66 Thus where Eimer went on to use his trends as
the basis of a major evolutionary mechanism, Cope tried to limit their
overall significance by insisting on the environmental factor. Cope does
not seem to have known of Eimer's later writings, since he never used the
term orthogenesis and actually coined his own name, physiogenesis, to cover
the direct production of variation by the external conditions.67 His work
thus illustrates how a neo-Lamarckian could subordinate the search for
regularity in evolution to environmental control, instead of using it as the
basis of internally directed orthogenesis.
The main problem with Cope's writing on variation was his uncritical
attitude toward the evidence he reported, whether from his own or from
other naturalists' studies. If a color variation seemed to be related to geo-
graphical distribution, it was automatically assumed that the organic change
was produced by the direct action of some factor in the environment.
Even where there was more definite evidence for an individual response to
the external conditions, it was again simply assumed that the variation
could be inherited and hence could serve as the basis of an evolutionary
mechanism.68 Such careless use of what often turned out to be irrelevant
evidence began to bring the neo-Lamarckian movement into disrepute,
especially after the rediscovery of Mendelism in 1900 reinforced the grow-
ing demand for experimental rigor in the study of heredity. Belief in the
possibility of directed variation did not die out overnight: Weismann him-
self had apparently confirmed the phenomenon on a small scale, and other
naturalists continued to look for hard evidence. But almost invariably the
examples that could be cited with any degree of plausibility were of an
extremely trivial nature, confirming Weismann's claim that the effect oc-
curred on such a small scale that it could never control the general course
of evolution.
At least one of the efforts to establish a more rigorous form of evidence
was associated with an attempt to present Eimer's theory in a more reason-
able light. In 1907 Vernon L. Kellogg, professor of entomology at Stan-
ford University, published his Darwinism todaya classic survey of the
66. Ibid., pp. 45-58. Much of this chapter is quoted directly from a popular article by J. A. Allen of
the U.S. Geological Survey.
67. Ibid., pp. 227-144. Physiogeneiij contrails with IrinetogcnesisCope's name for uic-inheritance
the latter being by far the more rignificant factor in evolution.
68. Ibid., pp. 230-237. The wont example is Cope's quotation from Edward B. Poulton, The
colours of animals: their meaning and use, especially considered in the case of insects (New York, 1890),
chapter vra, on insect coloration. Poulton was a prominent neo-Darwinian, and in the chapter fol-
lowing the one quoted by Cope he specifically noted that the color changes he had studied were not
hereditable; see Poulton, Colours of animals, p. 176.
anti-Darwinian arguments then in vogue. Kellogg was not a rabid op-
ponent of selectionism, indeed his careful analysis offers a good illustra-
tion of how the uncritical attitudes of writers such as Cope were exposed.
But neither was Kellogg completely satisfied with Darwinism, and he was
quite prepared to consider those alternatives that seemed to be supported
by sound evidence. In particular he accepted die existence of some non-
adaptive characters and the need to invoke a mechanism to explain them.
He discussed Eimer's work at some length, emphasizing that (unlike Na-
geli's theory) orthogenesis was scientifically respectable because it depended
on external forces to elicit the variation.69 A purely internal directing force
was unacceptable since it would provide no real explanation of why the
species was predisposed to vary in that particular way and would open the
way for the reintroduction of teleology. Kellogg passed over Eimer's in-
terpretation of the butterflies very quickly and does not seem to have
realized that in the end Eimer was using internal forces to impose an
exaggerated degree of unity upon the evolutionary process. Kellogg's own
line of evidence was of a far more restricted kind that merely suggested
die ability of a species to respond to a particular external influence. He had
found diat a certain species of beetle appeared to show a distinctively
marked form on the campus of Stanford University.70 Having vigorously
repudiated any utilitarian explanation of the strange markings, he was
forced to conclude that diey were the result of some form of directed
variation. Exactly why the campus beetles had been singled out in diis
way Kellogg was unable to say, but he almost certainly envisioned some
effect of the local conditions that had stimulated the change. Orthogenesis
dius became a small scale phenomenon that would probably be unique in
each case, not a series of extensive predetermined trends.
Kellogg and Weismann both de-emphasized the unifying aspect of or-
thogenesis as promoted by Eimer, but did so in two quite different ways.
Where Weismann postulated individual trends arising within die germ
itself, Kellogg allowed the germ to be modified in various directions by
the action of die external conditions. Both allowed orthogenesis, but only
at the expense of reducing it to a trivial level where it could never be ex-
pected to produce anything more dian the occasional nonadaptive charac-
ter. But for odiers it was precisely die regularity of orthogenesis, its poten-
tial for linking a series of species into a large scale trend, that deserved
69. Kellogg, Darwinism today (n. 1), pp. 280-285.

70. Kellogg, 'Ii there determinate variation?' Sdtna, 1906, N.s. 14,621-628. See abo Kellogg, Dar-
winism today (n. 1), p. 319.
attention. This was inevitable in paleontology, where a trend would only
become visible if it had acted consistently over a long period of time. But
Eimer's position also received occasional support from naturalists who
constructed their trends out of living species. Where the trends were per-
ceived to be large enough, they could not be dismissed as the result of
changing conditions, and this factor by itself threw the emphasis onto in-
ternal control as the only way of guaranteeing consistency.
The closest approach to Eimer's own position on the issue of internal
control of variation was made by James T. Cunningham, who had studied
at Oxford and later became naturalist to the Marine Biological Associa-
tion. In 1890 Cunningham translated Eimer's Organic evolution, proclaim-
ing in the introduction his own opposition to Weismann and support for
Lamarckism.71 He also noted a number of apparently nonadaptive char-
acters in various species that lent support to Eimer's concept of directed
variation. In 1896 Cunningham participated in a debate sparked off by a
defence of utility by Alfred Russel Wallace, the venerable co-discoverer of
natural selection who was still a staunch supporter of neo-Darwinism.
Cunningham challenged Wallace to demonstrate to him that any of the
characters normally used to distinguish one closely related species from
another had adaptive value.72 For anyone so strongly convinced of the
existence of nonadaptive characters, some form of orthogenesis would be
an essential ingredient of evolution.
Cunningham had speculated briefly on the nature of such a mechanism
in an article on the evolution offlatfishespublished in 1895. He postulated
an internal force producing nonadaptive variation, but did so in a way
that avoided Eimer's vision of an evolutionary process narrowly circum-
scribed by a unifying pattern. Cunningham specifically rejected genepis-
tasis and the claim that various modern species might represent stages in a
linear scale of development.73 There might be a tendency for related spe-
cies to vary in a similar way, but many of the broader connections that
Eimer had stressed could be explained by simple community of descent.
Evolution was essentially a branching, not a linear processperhaps one
of the most fundamental beliefs of the post-Darwinian era, but one which
71. Jama T. Cunningham, Translator'! preface,' in Eimer, Organic evolution (n. 8), ejp. pp. vii-xii.
72. Jamej T. Cunningham, The utility of ipecific characteristics,' Nature, 1896, 54, 295. Thii was
one of a series of response], mojtly critical, to A. R. Wallace, The problem of utility: are ipecific
characters always or generally useful?'_/. Linn. Sot. (zool.), 1896, 2J, 481-496.
73. James T. Cunningham, The origin and evolution of BztBsha,'Natural Science, 1895, 6,169-177,
233-239. P- 177-
Eimer had never fully come to grips with. But evolutionary divergence
was not solely due to the demands of adaptation: in addition there was
a tendency to definite variation, or growth in different directions, leading to
manifold variety of regular, definite symmetrical forms. This tendency can only
be regarded as internal to the organism, as connected widi the tendency to
growth and multiplication inherent in organic units.... Whatever the causes of
non-adaptive variation, the resulting structural features are the regular 'geomet-
rical' forms and characters which the multitude of different organic forms present
in such marvellous diversity.74
As a Lamarckian, Cunningham denied the independence of the germ from
its environment, and there was thus no reason why he could not have
anticipated Kellogg's suggestion that orthogenesis was also caused by ex-
ternal changes. But Cunningham, like Eimer, was fascinated by an element
of regularity that he perceived in certain organic relationshipsin this case
among different species offlatfishes.Such regularities among nonadaptive
features convinced him that variation must be directed by internal factors,
not by the caprice of the environment. On the other hand, Eimer had gone
too far in postulating trends governing the evolution of the whole animal
kingdom. For Cunningham, the character of each form of life determined
the possibilities of regular variation, and it was thus only in closely related
species that evidence of orthogenesis could be found.
Cunningham's belief that characters often have no adaptive value was
shared by many opponents of neo-Darwinism at the time. But his interest
in the regularity of some evolutionary developmentslimited as it wasis
more difficult to place. Theistic evolutionists such as William B. Carpenter
had sometimes used such a sense of regularity to argue for supernatural
direction of evolution. But although Cunningham's idea is not developed
in detail, he shows no sign of wishing to propose anything but a naturalistic
mechanism. In this Cunningham resembled Eimer himself, whose far
greater sense of the regularity of nature seems to have been derived from
the idealist world view of German Naturphilosophie. Probably Cunning-
ham derived his interest in the issue from his close study of Eimer's writ-
ings, while his more cautious approach reflected his less intimate connec-
tion with German thought. In fact, Cunningham does not seem to have
followed up his suggestion of 1895, and by 1900 he was involved in a
74. Ibid., p. 239. CimninghaTTi'i remarks on orthogcnesii attracted little attention, but his attempt
to confirm a Lamarckian explanation of the lick of coloration on the undenide offlatfishesas a direct
consequence of the absence of light was quite widely discussed.
quite different study of nonadaptive characters. In his Sexual dimorphism in
the animal kingdom he argued that secondary sexual characters are produced
by the direct stimulation of those parts of the body associated with repro-
ductive activity.75 Sexual dimorphism would thus result from variation
directed not by purely physical factorsinternal or externalbut by the
mating habits of the species. As such this study represented a notable shift
of interest away from Cunningham's earlier belief in regular orthogenesis,
since the sexual characters are generally unique to each species.
The limited extent of even Cunningham's involvement with regular
orthogenesis indicates the difficulties faced by anything resembling die
idealist viewpoint in post-Darwinian Britain. But the problem of the non-
adaptive characters themselves remained a thorn in the side of the neo-
Darwinians. Every effort was made to play down their significance, as in
the article by Wallace noted above. Some neo-Darwinians rejected the
whole idea of nonadaptive characters and mounted a campaign against the
efforts that were being made to explain animal coloration, for instance, in
a nonadaptive way. Edward B. Poulton, professor of zoology at Oxford
and a leading student of mimicry in the animal kingdom, argued that
mimicry and other forms of protective coloration gave clear evidence of
the close dependence of color schemes on adaptation. To explain mimetic
resemblances as die result of coincident variation due to internal forces, as
Eimer had done, was to ignore a multitude of factors proving that the
colors did have value for the protected species.76 But other neo-Darwin-
ians were more responsive to the widespread claim that some characters
seem to have no utilitarian value. We have seen that Weismann himself
eventually used germinal selection to allow for die production of such
characters, and he was actually anticipated in diis application of his mech-
anism by Alfred Russel Wallace. Since Wallace soon became unhappy
with Darwin's dieory of sexual selection, he was disturbed by the exu-
berance of some secondary sexual characters. Wallace could see how such
characters began as recognition marks, but he thought that utility could
not explain how they grew to die extent observed, for instance in the
plumage of some birds. In a letter to Raphael Meldola in 1896, Wallace
confessed this difficulty and argued for germinal selection as the solution
75. Jama T. Cunningham, Sexual dimorphism in the animal kingdom: a theory of the origin of secondary
sexual characters (London, 1900).
76. Edward B. Poulton, 'Natural idection and the cause of mimetic resemblances and common
warning coloun" (1898), reprinted in Poulton, Essays on evolution, 1889-1907 (Oxford, 1908), pp. 220-
270. On Eimer ice p. 224. See also Poulton, Colours of animals (n. 68).
77
to it. A character could be primed for development by utility, but ger-
minal selection would then be able to increase it up to any size not posi-
tively harmful to the species. For Wallace, at least, the possibility of varia-
tion directed by germinal selection solved a real problem created by the
existence of apparently useless characters.
Although there were some positive reactions to germinal selection, the
mechanism did not become widely popular. The attention of most natu-
ralists was distracted by the rediscovery of Mendelism in 1900 and the
subsequent publication of Hugo De Vries' mutation theory. As late as 1908
one of Weismann's leading British supporters, J. Arthur Thomson, was
still treating both Mendelism and germinal selection as equally important
extensions of the original germ plasm theory.78 But in fact the extensions
were in opposite directions, and Mendelism was often seen as hostile to
neo-Darwinism. Mendel's theory postulated the existence of germinal
units, while the mutation theory supposed that these units could undergo
abrupt changes from time to time. For the Mendelian the germ was made
up of fixed units; it was not an active system in which one element could
benefit at the expense of others, as in germinal selection. In addition both
Mendelism and the mutation theory emphasized the discontinuity of vari-
ation, and to begin with it was widely assumed that such discontinuity
must undermine the neo-Darwinians' insistence on continuous evolution.79
The mutation theory offered a new way to explain nonadaptive charac-
ters without the necessity of directed variation. De Vries insisted that the
selection of small individual differences was incapable of producing per-
manent change in a species. New varieties and even species were formed
directly by sudden mutation, and since no selection was involved the new
characters need not have adaptive value. The mutations occurred in ran-
dom directions and in De Vries' opinion were themselves subject to com-
petition and selection. Nonadaptive features could exist only in the short
77. Wallace to Meldola, 12 October 1896, in James Marchant, ed, Alfred Russel Wallace: le ten
and reminiscences (New York and London, 1916), pp. 320-331. Germinal selection wai also praised by
F. A. Dixey, a colleague of Poulton at Oxford: Nature, 1896, 54, 121-112. But Dixey shared Weis-
mann'i original view that the real value of germinal election wai in tolving the problem of the
disappearance of useless characters, not their production. The fact that there ii no mention of germinal
selection in Poulton'j own Essays (n. 76) suggests that he did not share Dixey'1 interest in this par-
ticular aspect of Wcismann's work.
78. J. Arthur Thomson, Heredity (New York and London, 1908); on germinal selection see pp.
458-475, on Mendelism chapter x.
79. On the debate between the neo-Darwinians and Mendelians, and their subsequent reconciliation
via the genen'cal theory of natural selection, see William B. Provine, The origins oftheoretical population
genetics (Chicago, 1971).
term, since the broad course of evolution was still determined by selection
and utility. De Vries examined and rejeaed the possibility that there might
be a tendency for series of mutations to occur consistently in the same
direction.80 However, the leading American supporter of the mutation
theory, Thomas Hunt Morgan, realized that mutation itself might be able
to direct the course of evolution. In 1903 in his Evolution and adaptation,
Morgan strongly opposed the utilitarian foundations of Darwinism, sug-
gesting that any mutated form not hopelessly incompatible with the en-
vironment would be able to establish itself.81 Believing that many well-
developed characters have no adaptive value, Morgan postulated that in
some cases a series of related mutations might push evolution in the same
direction, thereby in effect producing orthogenesis.82 While opposing Na-
geli's vision of evolution driven completely by an internal force, Morgan
nevertheless saw the possibility that germinal factors might sometimes im-
pose a direction upon the sequence of mutations. He admitted, however,
that he had no real evidence for such a phenomenon, and eventually aban-
doned it along with De Vries' theory of evolution by sudden leaps. By
1916 Morgan had completely reversed his attitude toward utility and se-
lection, largely as a result of his work on true genetic mutations in the
fruit fly, Drosophila. He then began to argue for the selection of individual
differences introduced into the population by small mutationsan antici-
pation of what was to become the genetical dieory of natural selection.83
The question of whether variation was totally random or sometimes
directed along fixed lines was thus debated within both the rival neo-Dar-
winian and mutationist camps. The common belief in the widespread exis-
tence of nonadaptive characters had in both cases provided support for the
possibility of orthogenesis. But Morgan's gradual acceptance of selection
and utility symbolized the collapse of this position within most areas of
natural history except paleontology. The final development of the genet-
ical theory of natural selection through the statistical techniques of R. A.
Fisher and others in the 1930s re-established the need for adaptation as the
driving force of evolution and proclaimed the need for largely random
80. Hugo De Vries, The mutation theory: experiments and observations on the origin of spedcs in the
vegetable kingdom, trim. J. B. Farmer and A. D. Darbyshire, 2 vols. (London, 1910), 1,66. See Garland
E. Allen, 'Hugo De Vries and the reception of the "mutation theory," ' J. Hist. BioL, 1968, 113-
139 and Peter Bowler, 'Hugo De Vriej and Thomai Hunt Morgan: the mutation theory and the spirit
of Darwinism,' Ann. Sd. 1978, 35, 55-73.
81. Thomas Hunt Morgan, Evolution and adaptation (1903; reprint ed, New York, 1908), pp. 348,
464.
8a. Ibid., pp. 292-293, 461.
83. Thomas Hunt Morgan, A critique of the theory of evolution (Princeton, 1916).
mutation as a source of the variations upon which selection must act.
Within this new synthesis there was only one possible way in which vari-
ation itself could influence the course of evolution: not as a positive direc-
tional influence but as a negative factor defining limits within which selec-
tion had to work. An early version of this more sophisticated concept of
directed variation had already been proposed at the turn of the century by
the leading American naturalist Charles Otis Whitman.
As professor of zoology at the University of Chicago and first director
of the Marine Biological Laboratory at Woods Hole, Whitman was a
powerful if somewhat unorthodox supporter of neo-Darwinism. His in-
terest in embryology and his sense of the complexity of individual devel-
opment led Whitman to reject both Mendelism and the mutation theory
with their neat simplifications of unit characters and single evolutionary
steps. Whitman preferred the Darwinian alternative of selection acting
upon small variations, but a detailed study of color variation in pigeons
led him to doubt that the variations occurred in all possible directions.
This work was performed between 1900 and 1910, although its publication
was only completed posthumously in 191 p. 84 Whitman found that there
was a definite tendency for variations to occur more readily in some di-
rections than in others. He compared various races of pigeons to show that
(in contrast to Darwin's opinion) their tendency was always to move from
a checkered to a barred pattern.85 Since there was no guarantee that a
systematic arrangement of the modern forms corresponded to the actual
course of evolution, Whitman supported his view with an appeal to the
recapitulation theory and a study of the development of individual birds.86
Further research suggested that a similar tendency to vary had influenced
the evolution of other groups of birds.87
Whitman read Eimer's accounts of orthogenesis very carefully and re-
alized that Eimer's approach tended to emphasize rigid control of variation
84. Charles Otis Whitman, Posthumous works, 3 vols., ed. Oscar Riddle (Washington, 1919), 1,9-36.
85. Ibid., pp. 15-36. Further details are given in chapter 3, pp. 37-48, originally written in 1905 as
a continuation of The problem of the origin of species' but not previously published.
86. Ibid., chapter 4, "The origin and relationship of the rock-pigeons as revealed in their color pat-
tern,' pp. 49-63, see p. 51. This chapter was originally an address read to the National Academy of
Sciences in 1903, of which only an abstract had been published. For Whitman's general views on the
recapitulation theory see p. 7 and chapter 10, 'The problem of organic developmentfacts and theo-
ries,' pp. 177-194. The latter was originally an address given to the 7th International Congress of
Zoology, Boston, 1907, not previously published.
87. Ibid., chapter 6, "The turtle dove pattern in other orders of birds,' pp. 117-139. Note that in
this chapter Whitman specifically contradicts some of Eimer's 'laws' of orthogenesis, see pp. 118120
and p. 134.
by internal factors.88 Most Darwinians were suspicious of the internal
control of variation because such absolute predetermination of evolution
tended to reintroduce the problem of teleology. Whitman argued that a
more limited kind of orthogenesis might be postulated without implying
that the trends were goal-directed. If the existing germinal factors respon-
sible for individual growth possessed a certain structure that allowed
change to occur more easily in some directions than others, the process
would require teleology only in the same sense as embryology itself. Whit-
man wrote:
I take exception here only to the implication that a definite variation-tendency
must be considered to be teleological because it is not 'orderless.' I venture to
assert that variation is sometimes orderly and at other times rather disorderly,
and diat the one is just asfreefromteleology as die odier. In our aversion to the
old teleology, so effectually banished from science by Darwin, we should not
forget diat die world is full of order, die organic no less dian die inorganic.
Indeed, what is die whole development of an organism if not stricdy and mar-
velously orderly? Is not every stage, from die primordial germ onward, and die
whole sequence of stages,rigidlyorthogenetic? If variations are deviations in die
directions of die developmental processes, what wonder is mere if in some di-
rections dicre is less resistance to variation than in odiers? What wonder if die
organism is so balanced as to permit of bodi unifarious and multifarious varia-
tions? If a developmental process may run on diroughout life (e.g. die lifelong
multiplication of die surface pores on die lateral-line system in Arnia), what
wonder if we find a whole species gravitating slowly in one or a few directions?
And if we find large groups of species all affected by a like variation, moving in
die same general direction, are we compelled to regard such a 'definite variation-
tendency' as teleological, and hence out of die pale of science? If a designer sets
limits to variation in order to reach a definite end, die direction of events is
teleological; but if organization and the laws of development exclude some lines
of variation and favor odiers, there is certainly nodiing supernatural in diis, and
nothing which is incompatible widi natural selection. Natural selection may
enter at any stage of orthogenetic variation, preserve and modify in various
directions die results over which it may have had no previous control.89
Whitman was unable to give details of how the existing developmental
process imposed restrictions on the range of possible variation. His work
could not be incorporated into the new genetics and was ignored during
the creation of die genetical theory of selection, where for simplicity it
88. Ibid,, p. 9. whitman criticizes Eimcr for hii 'intemperate ferocity' toward Darwinism and ar-
gues that his rejection of the utilitarian viewpoint automatically tended toreintroduceteleology.
89. Ibid., p. 11.
was usually assumed that variation is random. But more recently it has
been recognized that the existing genotype does indeed impose some limit
on the range of possible variations, and Ernst Mayr has hailed Whitman's
theory as an important anticipation of the modern view.90 In fact Whit-
man appears to have granted orthogenesis a rather more positive role than
would be admitted today. But Mayr's point is valid in the important sense
that Whitman was suggesting restrictions on variation rather than the ab-
solute predetermination of future events.
The genetical theory of selection did not sweep the board immediately
in 1930, since occasional efforts were still made to postulate a far more
positive control of variation than Whitman's approach permitted. Richard
Goldschmidt, for instance, possessed an even deeper sense of the com-
plexity of individual growth than had Whitman himself, a sense which led
him to suggest that in many cases there was only one possible mutation
that would allow the species to make a viable change.91 But Goldschmidt
also attributed evolution to macro-mutations, and along widi his unortho-
dox views on genetics this ensured that his ideas did not gain wide accep-
tance. His work represents perhaps the last effort to promote a theory of
rigidly directed variation to appear outside the area of paleontology. The
genetical theory of selection effectively synthesized the new genetics and
the old Darwinian natural history, and in so doing re-established the prin-
ciple of utility to the satisfaction of most biologists. Not only had it be-
come difficult to conceive of genetic mechanisms that would produce non-
adaptive trends, but studies in the field were increasingly throwing the
opponents of useful adaptations onto the defensive by confirming the
adaptive value of most characters.92
In the circumstances, it is hardly surprising that paleontologists provided
the last line of defense for orthogenesis. Of all biologists, they were the
least directly concerned with the actual mechanism of variation and the
least likely to take the new genetics seriously. At die same time their field
offered little or no opportunity to study the potential utility of any unusual
character. In 1912 die American paleontologist Henry Fairfield Osborn,
perhaps the leading twentieth century supporter of orthogenesis, consid-
9a Dictionary of scientific biography, t v 'Whitman, Charles Otis.' On the topic ofrestrictedvaria-

tion see Huxley, Evolution (n. 3), pp. 515-524.
91. Richard Goldschmidt, 'Some aspectt of evolution,' Science, 1933, 78, 530-547, p. 544. Alio
Goldschmidt, The material basis of evolution (1940; reprint ed., Patterson, N.J., i960), p. 331. See
Garland E. Allen, 'Opposition to the Mendelian-chromoiome theory: the physiological and develop-
mental genetics of Richard Goldschmidt,' J. Hist. Biol, 1974, 7, 49-92.
92. On the debate over utility in the 1930s sec Julian Huxley, Evolution (n. 3), p. 417.
ered the problems facing those who were trying to construct a mechanism
for evolution and concluded that no satisfactory solution was in sight.
Osborn thought that biologists might have to remain satisfied with abstract
laws of development that merely described the findings of the paleontolo-
gist.93 For this he was accused by Thomas Hunt Morgan of toying with
mysticism, and Morgan's work on genetics was already helping to lay the
foundations of what would become the genetical selection theory.94 Yet
in the 1920s and 30s, Osborn continued to advocate orthogenesis, while
confessing his inability to provide a mechanism that would explain how a
trend of variation could become fixed in the germ plasm.95 Osborn's
studies of characters such as the horns and teeth of the Titanotheres re-
vealed evolutionary trends that appeared to begin with no adaptive value,
then moved into an adaptive phase, but finally ran on to overdevelopment
and extinction. Osborn's work thus provided not only the last line of sup-
port for orthogenesis but also the last apparently convincing demonstra-
tions of the existence of nonadaptive characters. Even the supporters of the
genetical theory of selection accepted that in such cases the overdevelop-
ment would have to be explained as a function of allometry (differential
growth-rates) radier than adaptation.96 Only more recently has it been
argued that even the most bizarre structures originally attributed to over-
development might have had a real value to the species.97
CONCLUSION
The rise and fall of orthogenesis represents an important component of

the debate over the validity of Darwin's theory which raged around the
turn of the century. As one of the most obvious mechanisms for the pro-
duction of what were widely thought to be nonadaptive characters, ortho-
genesis challenged not only selection but also the principle of utility itself.
Weismann's retreat into germinal selection suggests that the problem of
directed variation was perceived as a major threat by the neo-Darwinians,
and only with the development of the far more sophisticated genetical
theory of natural selection did the enthusiasm for orthogenesis and the
93. Henry Fairfield Osborn, The origin and evolution of life (New York, 1913), p. xi.
94. Morgan to Osbom, 26 December 1917, quoted in Garland E. Allen, "Thomas Hunt Morgan
and the emergence of a new American biology,' Quart. Rev. Biol., 1969,44,168-188, lee pp. 179-180.
95. Henry Fairfield Osborn, The Titanothats of ancient Wyoming, Dakota and Nebraska, a vols.
(Washington, 1919), n, 849.
96. Julian Huxley, 'Constant differential growth-rates and their significance,' Nature, 1924, 114,
895-896 and A. H. Sturtevant, 'An interpretation of orthogenesis,' Science, 1924, $g, 579-580. See also
Julian Huxley, Problems of relative growth (London, 1932), especially pp. 214-221.
97. See for instance Gould, 'Origin and function' (n. 60).
opposition to utility begin to die away. Yet the basic idea of orthogenesis
could be exploited in different ways, each of which created its own prob-
lems. Given the existence of nonadaptive variation-trends, biologists had
to decide whether the chief controlling influence was derived externally
from the environment or provided internally by the existing constitution
of the organism. Acceptance of internal control posed the particular prob-
lem of explaining how such a predetermined pattern of evolution might
be invoked without assuming teleology.
The most obvious scientific question involvedwhether or not the
germ plasm was isolated from its surroundingsdid not dictate the posi-
tions taken up by scientists on the issue of external or internal control of
orthogenesis. Those who believed that the germ could be influenced from
outside were free to claim that variation was directed by environmental
factors. This was the position taken up superficially by Eimer and more
seriously by Kellogg. Yet Eimer's efforts to expand the scope of ortho-
genesis led inevitably to the conclusion that there must be a strong element
of internal control. Only by internal control could trends be produced on
a scale large enough to serve as major unifying factors within the pattern
of evolution. Weismann was also forced to explore the possibility of in-
ternal control by the very nature of his germ plasm theoryyet as a Dar-
winian his main concern had to be the elimination of the element of regu-
larity that so fascinated Eimer. Germinal selection was a theory of internally
controlled variation which imposed no fixed pattern on the directions that
might arise. It proved impossible to formulate the equivalent of such a
mechanism within the new genetics, and eventually the revival of utility
made the search for positive internal control unnecessary. All that was left
was Whitman's much more flexible concept of limits imposed on the
variations among which natural selection might choose.
If the debate over the nature of heredity did not dictate attitudes toward
orthogenesis, we must look elsewhere to understand the scientists' moti-
vations. Why did so many naturalists try to emphasize the prevalence of
nonadaptive characters? Why were they sometimes tempted to arrange
either living or fossil species into neat linear sequences advancing in parallel
along the same predetermined path? Loren Eiseley has explained the gen-
eral opposition to utility as the reflection of a growing moral disapproval
of the Darwinian picture of a world driven by struggle, and of the harsh
social policies sometimes based on this biological model.98 There seems
98. Eiseley, Darwin's century (n. 1), pp. 250-251. Eiseley isreferringto De Vries' mutation theory,
but the same point applies to orthogenesis.
little doubt that Eimer himself found the Darwinian world view unac-
ceptable for broadly philosophical reasons, and the same must be true to
some extent of his followers. Yet the simple existence of nonadaptive char-
acters could be accounted for in a number of ways, including both ortho-
genesis and De Vries' mutation theory. The choice between pure muta-
tionism and orthogenesis would be conditioned by the scientist's attitude
toward the most fundamental difference between them. As De Vries pre-
sented it, the mutation theory still demanded that evolution be the result
of largely random variation, while the whole purpose of the more extreme
versions of orthogenesis was to stress the orderliness of development. Those
who wished to restore to the evolutionary process the element of regu-
larity denied by the materialism of Darwin's theory would inevitably tend
to opt for orthogenesis. In Eimer's case, the expectation that nature ought
to exhibit a regular pattern of development seems to represent the last
phase in the influence of the old idealist world view of Naturphibsophie.
Finally, to what extent did the search for order in nature really threaten
the mechanistic interpretation of science that had been promoted by Dar-
winism? By the end of the nineteenth century, most biologists expressed a
determination to avoid the perils of mysticism or a reintroduction of the
supernatural. Nageli's 'perfecting principle' was often cited as a clear ex-
ample of what had to be avoided. Thus an isolated attempt by a German
scientist to revive such a principle was dismissed by Kellogg as 'opposed,
in sharpest contrast, to the very spirit of science.'99 Eimer himself made
strenuous efforts to dissociate his theory from Nageli's, claiming that the
degeneration sometimes brought about by orthogenesis indicated that the
trends were not goal-directed. Yet Eimer left many questions unanswered
when he postulated vast unifying trends in evolution for which he could
give no truly natural explanation. It was no coincidence that Henri Berg-
son's Creative evolution referred to the work of both Theodor Eimer and
Edward Drinker Cope to support the claim that evolution was guided and
unified by the mysterious Man vital.100 Whatever the intentions of the
biologists concerned, the temptation to exaggerate the scope of orthogen-
esis inevitably compromised the materialist interpretation of evolution.
Darwin's theory not only eliminated teleology, it also undermined the
belief that the structure of nature ought to display regular patterns at any-
thing beyond the microscopic level. The laws of nature are regular, but
99. Kellogg, Darwinism today (n. l), p. 278, following Ludwig Plate's opinion of Georg Pfcfler'i
Die Entwickclung of 189J.
100. Henri Bergion, Creative evolution, tram. Arthur Mitchell (New York, 1911), pp. 73-77.
the whole system is so complex that the interaction of natural laws may
not be expected to generate neat patterns in the broad structure of the
world we observe. As a reflection of the belief that there were such orderly
patterns in the evolution of life, the more extreme versions of orthogenesis
were almost impossible to reconcile with the materialist philosophy.
Department of History
University of Winnipeg

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Theodor Eimer and Orthogenesis:

Evolution by 'Definitely Directed Variation'

I U M E R O U S objections were raised against Darwin's

Although Darwin had realized that variation is normally random, he knew

12. Ibid., pp. 134-139.

43. Ibid. (n. a), p. 33; (n. 30), p. 38.

In 1888 Eimer proclaimed his opposition to Weismann's neo-Darwinism

ji. Webmann, Theory of descent (n. 46), 1, 363.

57. Weismann (n. 10), pp. 275-276.

In 1897 Eimer published a vigorous critique of germinal selection.62 In his

69. Kellogg, Darwinism today (n. 1), pp. 280-285.

9a Dictionary of scientific biography, t v 'Whitman, Charles Otis.' On the topic ofrestrictedvaria-

The rise and fall of orthogenesis represents an important component of

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