Drosophila. A Life in The Laboratory. Kohler
Drosophila. A Life in The Laboratory. Kohler
ROBERT E. KOHLER
INTRODUCTION
Few laboratory creatures have had such a spectacularly successful and productive history as Drosophila, the friendly fruit
fly. It first entered laboratories about 1901, revealed its talent for
experimental genetics in 1910, and with some ups and downs is still
going strong. If not the first "standard" laboratory creature,
Drosophila is certainly representative of the type. It has been a
standard laboratory animal for so long that we no longer think much
about how, as a wild creature, it has brought into an alien environment and transformed into an instrument of scientific production.
Why did it become such a fecund and cosmopolitan species of
laboratory animal? What was it in the constitution of the little fly
and in the social practices of experimental geneticists that made
them such productive symbionts? These questions can be addressed
in many ways: as the history of ideas, for example, or as institutional history. Here, however, I shall treat the question of
introduction ecologically, as natural history.
It is a commonplace of ecological history that landscapes, which
may seem to be "natural," were in fact fashioned by humankind's
technologies of fire, plow, and seed. 1 But no less can domestic
1. See, for example, Carl Sauer, "The Barrens of Kentucky," in Land and Life:
A Selection from the Writings of Carl Ortwin Sauer, ed. John Leighly (Berkeley:
University of California Press, 1969), pp. 23-31 (first published in 1927); Carl
Sauer, "Man's Dominance by Use of Fire," in Selected Essays 1963-1975
(Berkeley: Turtle Island Foundation, 1981), pp. 129-156; William Cronon, Changes
in the Land: lndians, Colonists, and the Ecology of New England (New York:
Hill and Wang, 1983); Richard White, The Roots of Dependency: Subsistence,
Environment, and Social Change among the Choctaws, Pawnees, and Navajos
(Lincoln, Neb.: University of Nebraska Press, 1983); and Alfred Crosby, Jr.,
Ecological Imperialism: The Biological Expansion of Europe, 900-1900
(Cambridge: Cambridge University Press, 1986).
The following abbreviations are used below to cite archival sources: AHS
(Alfred H. Sturtevant Papers, Archives, California Institute of Technology,
Journal of the History of Biology, vol. 26, no. 2 (Summer 1993), pp. 281-310.
9 1993 Kluwer Academic Publishers. Printed in the Netherlands.
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and change the purposes for which they had originally been brought
into the laboratory. They drove out other favorite creatures of
experimental biologists. We need to see the relation between
Drosophila and drosophilists ecologically, as an interactive and
evolving symbiosis.
What, then, was Drosophila before it entered the laboratory?
How did it cross that ecological threshold, and how, having gained
a niche in a new ecosystem, did it establish itself as a dominant
species, upon which the fly people came to depend absolutely for
their professional livelihoods? The natural history of the fly is not
quite as rational as it is sometimes portrayed. Drosophila was
brought into the laboratory not to do genetics but because it seemed
promising for studies of experimental evolution, and because its
natural cycles and habits fit the natural history of academic life.
Its capacity for genetic work was revealed unexpectedly.
THE NATURAL HISTORY OF DROSOPHILA
The genus Drosophila of the family Drosophilidae is made up
of more than nine hundred species, which inhabit every corner of
the world except the driest deserts and the coldest tundras.
Drosophilas need moisture (their name means "dew lover"), a
temperate climate, and yeasts, which is what they feed on. ("Fruit
fly" is thus a misnomer; "yeast fly" would be more apt.) Where
there is decaying vegetation or fruit to support colonies of yeasts,
there are fruit flies. Some species are highly local and specialized, feeding exclusively and shyly around forest fungi, flower
pollen, water plants, fruits, or tree sap; such specialists tend to be
rare, however, and are seldom seen except by field naturalists. Other
species are less finicky eaters and forage widely and opportunistically; they get around and are more ubiquitous and visible.
Specialization is the rule: about 90 percent of all Drosophila species
are endemic to just one of the six great ecological regions of the
world; just eight species groups inhabit all six regions. 4 These
"cosmopolitan" groups were the ones that found their way into
biologists' laboratories. They were the only ones in a position to
do it; experimental biologists had no reason to venture from their
laboratories into forests to seek out rare species and bring them
back alive. It was the cosmopolitan species that were all about,
4. John T. Patterson and Wilson S. Stone, Evolution in the Genus Drosophila
(New York: Macmillan, 1952), pp. 6-7, 48; Alfred H. Sturtevant, The Classification
of the Genus Drosophila, with Description of Nine New Species (Austin: University
of Texas, publication no. 4213, 1942), pp. 5-51.
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just on the other side of the threshold of the laboratory. Just leave
a ripe banana on a window sill for awhile, put it in a jar, and wait
for the drosophilas to hatch: chances are you will have D.
melanogaster, the standard laboratory fly; you can tell it by the
black bottom of the males (hence its name).
The cosmopolitan species of Drosophila are seasoned and hardy
travelers. Blown about by wind and storm, they have been collected
from airplanes 3000 feet in the air - but mainly they have traveled
with humankind, of whom they are devoted hangers-on, especially
D. melanogaster. The distribution of this species group suggests
that it evolved in Southeast Asia and then spread around the world.
It probably got to the New World from the Mediterranean and North
Africa via slave ships, and from the Caribbean it traveled north
along the trade routes of the international traffic in rum and sugar
and the new trade in bananas and fresh fruits that burgeoned after
the American Civil War. It is no accident that D. melanogaster,
or D. ampelophiIa as it was first called, was first sighted in the
eastern cities of North America in the 1870s. 5 Melanogaster is the
great hitchhiker among the drosophilas, as Homo sapiens is the
great traveler among the primates; wherever its human symbionts
have migrated, D. melanogaster has gone along for the ride.
It was agriculture and fruit growing that made possible this symbiosis between drosophilas and humankind - agriculture, and its
associated preserving and fermenting technologies. This association is clearly revealed in drosophila's common names: vinegar
fly, wine fly, pickled fruit fly, pomace fly (pomace is the apple mash
left over from cider pressing). "Ampelophila" means lover of vines
- that is, grape vines. Drosophilas, especially fruit feeders like D.
melanogaster, are so attached to humankind because their hosts
are such copious consumers of fermented fruits and such abundant
producers of organic waste and garbage. Drosophitas flourish in
vineyards and orchards, breweries, cider mills, pickle works. They
breed prodigiously in the piles of discarded and rotting fruits and
vegetables that surround citrus orchards and fruit wharves. In the
cold of North American cities and towns drosophilas have wintered
over in fruit and root cellars (in the days before refrigeration), or
in fruit stores and restaurants. Most of the cosmopolitan species are
fruit feeders, in fact, and use the north-south trade in tropical fruits
to reoccupy the north each year, congregating wherever fruits
accumulate at centers of production, distribution, processing, and
5. Patterson and Stone, Evolution, pp. 52-55; T. H. Morgan, Calvin B. Bridges,
and Alfred H. Sturtevant, "Genetics of Drosophila," Bibliog. Genet., 2 (1925),
5-6.
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disposal. In cities and suburbs hardier species of these inconspicuous stowaways and hitchhikers thrive around garbage pails and on
municipal dumps. 6 These cosmopolitan species of Drosophilawere
founding partners in the complex ecological system of human agriculture, along with domesticated plants and animals, pests, and
parasites.
Drosophilas were not domesticated creatures, obviously, but
commensals that had adapted to the domestic economy of
humankind. It is no accident that experimental biologists eventually took up with Drosophila, nor that the cosmopolitan D.
melanogasterbecame the standard laboratory fly: of all the species
of Drosophila it was the most likely to be associated with urban
academic biologists, the most at home among concentrated
humanity. It was the most accustomed to crossing the threshold
between indoors and out, the most opportunistic and versatile
forager, and thus the most likely to find its way into a banana-baited
trap and an experiment.
EXPERIMENTAL BIOLOGISTS AND NEW ORGANISMS
For Drosophila to become a domesticated creature it had first
to enter the laboratory. It had to venture from its natural home in
the gardens and dustbins of humankind into the domain of the
creators of a second, experimental nature. The question is, when
and why was Drosophila brought indoors - not accidentally, as a
pest to be brushed aside or squashed, but deliberately, as a fellow
laborer to be disciplined and worked?
As Garland Allen has show, T. H. Morgan was one of half a
dozen or so people who began to employ Drosophila for experimental work in the early 1900s. The first to use them were William
E. Castle and his students at Harvard University, starting in 1901,
and most other early drosophilists got the idea directly or indirectly
from them. William J. Moenkhaus brought Drosophila into his
laboratory at the Indiana University Medical School in 1903, having
learned of them from Castle. Around 1905 Moenkhaus persuaded
the entomologist Frank E. Lutz that Drosophila might be a good
organism for his work on experimental evolution at the Carnegie
Institution's Station for Experimental Evolution at Cold Spring
Harbor, Long Island. It was probably Lutz who introduced Morgan
to Drosophila about 1906, and in 1907 Morgan passed it along to
6. Patterson and Stone, Evolution, pp. 91-92; Warren P. Spencer, "Collection
and Laboratory Culture," in Biology of Drosophila, ed. Milislav Demerec (New
York: Wiley, 1950), pp. 547-549.
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aphids with a usefully odd sex life. For Mendelizing, the different
varieties of domesticated and wild mice and rats seemed the best
bet, as did insects like Drosophila for studies of mutation and
evolution. And those were just the major lines. Morgan also
centrifuged eggs of mollusks, hoping to get aberrant distributions
of chromosomes, and tried (in vain) to find a reliable source of
Western song sparrows, which he hoped to resettle in the Bronx
Zoo for a study of adaptive variation. When a graduate student
was told by his doctor to find outdoor work, Morgan got the idea
of sending him to Hawaii to collect the multitudinous species of
land snail, which had proliferated there in isolated coastal valleys,
for a study of variation and speciation in the wild. He kept a small
colony of pigeons for breeding work (and later guinea pigs and
fowl, too) and avidly attended fancy dog and poultry shows. A.
H. Sturtevant reckoned that Morgan worked with over fifty different
species, n "That man has more irons in the fire," Frank Lutz
quipped, "than an ordinary man has coals. ''12 It was true; Morgan
himself poked fun at his habit: "I am doing many things as usual,"
he told his friend Hans Driesch, "also as badly. ''13 But as a foraging
strategy it paid off.
Other experimentalists also foraged widely, but usually within
a single kingdom of the natural or seminatural worlds. In 1910
Castle reported that he maintained colonies of 400 rabbits, 700
guinea pigs, 500 mice, 1000 rats, 400 pigeons, and 8 dogs. He
had also had some yellow cats, but they had "up and died," and
the frogs did not count. 14 Frank Lutz was similarly far-ranging
within the world of insects, working on Gryllus (crickets) and, as
time allowed, Crioceris, Hyphantia, and Spilosoma. 15 Morgan's
eclecticism was somewhat extreme, but it exemplified a generational style.
Morgan's habit of trying out new organisms was a conscious
strategy for keeping up with or insinuating himself into fast-moving
research fronts. He was a great reader and forager of the litera11. A.H. Sturtevant, "Thomas Hunt Morgan," Biog. Mem. Nat. Acad. Sci.,
33 (1959), 296-298; Morgan to C. B. Davenport, April 30, 1904; July 22, 1905;
June 25, July 4, 1906 (sparrows); Morgan to Davenport, January 23 and [c. 27],
1905 (snails); Morgan to Davenport, June 5, 1905; February 5, 1907 (shows), all
in CBD.
12. F. Lutz to C. Zeleny, October 30, 1908, CZ 2.
13. T.H. Morgan to H. Driesch, October 23, 1905; also September 15, 1907,
both in THM-APS; Morgan to E. G. Conklin, July 2, 1904, EGC.
14. W.E. Castle, "Report," September 1, 1910, CBD.
15. Frank B. Lutz, "Breeding Strains of Insects," Carnegie Inst. Wash. Yearbk.,
6 (1907), 78.
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Drosophila's crossing of the threshold between field and labo16. T.H. Morgan to H. Driesch, January 30, 1909, THM-APS.
17. T.H. Morgan to C. B. Davenport, June 5, 1905; December 5, 9, 12, 14,
24, 1905; January 24, April 5, August 20, 1906; April 15, 1907, all in CBD;
Morgan to H. Driesch, February 23, August 16, September 15, 1906; November
27, 1907; January 20, 1908, all in THM-APS; Allen, Morgan (above, n. 8), pp.
132-136.
18. T. H. Morgan to C. Zeleny, September 30, November 11, 12, 1906; May
2, 1907, all in CZ; Morgan to C. B. Davenport, July 4, 1906, CBD.
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7).
22. W. Moenkhaus, "The Effects of Inbreeding and Selection on Fertility,
Vigor and Sex Ratio of Drosophila ampelophila," J. Morph., 22 (1911), 123-154.
23. F. Payne, "Forty-nine Generations in the Dark,", Biol. Bull., 18 (1910),
188-190; F. Payne to C. Zeleny, October 7, 1907, CZ 2.
24. Lutz, "Breeding Strains" (above, n. 15).
25. F. Lutz to C. Zeleny, February 24, 1907, CZ 2; Lutz to C. B. Davenport,
August 3, 1908, CBD.
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Drosophila: A
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filled with rotting fruits and vegetables. Although their populations soared and crashed with the seasons, they were accustomed
to wintering over in human habitations and happily bred year-round
in the warmth of laboratories. They were not fastidious feeders
and thrived on anything that would support a growth of yeast;
bananas quickly emerged as the ideal, and the international trade
in tropical fruits guaranteed a year-round supply of cheap fly food.
Best of all, drosophilas were free and easily replaced - so if colonies
were killed off by inexperienced or careless students, only a modest
investment was lost. When cultures died out in the summer vacation
it was easy to start new cultures when student laborers returned
in the fall. The annual cycle of drosophilas and students coincided. The craft skills of Drosophila culture were easily learned,
and a fly colony could be easily maintained in the interstices of a
busy undergraduate schedule. Animal colonies and greenhouse collections were far more exacting o f time and skill, and were too
valuable to entrust to succeeding generations of green apprentices.
Also, drosophilas were inconspicuous and innocuous - unlike, say,
rodents. Castle was called on the carpet in 1903 to reassure Harvard
President Charles W. Eliot that his hordes of rats and mice would
not escape from their improvised quarters and devour books
belonging to other occupants of their shared space. 3~
Drosophilas were no less useful to professors who wished to
demonstrate biological principles to their classes with living
material. Whereas many insects did not winter over, and animals
were expensive, Drosophila was ideal for short-term student
projects on tropisms, sexual dimorphism, metamorphosis, and so
on. Lutz sung the praises of the little fly to biology teachers - "a
most excellent laboratory material. ''31 Drosophila, in short, was
nicely adapted to the collegiate environment and the cycle of
academic seasons. And so it was that fruit flies gained a small niche
among their more cultivated and prestigious competitors in the
second nature of academic laboratories.
Drosophilas, with their low status and undemanding lifestyle,
were well adapted to the domestic economy of collegiate and university research. Cash poor but rich in cheap, eager, and renewable
student labor, departments of biology made the most of what they
had. Student scholarships and assistantships were a major subsidy
30. W. E. Castle to Jerome Greene, January 1, 1903; Greene to Castle,
December 29, 1902, both in Charles W. Eliot Papers 133/897, Pusey Library,
Harvard University, Cambridge, Mass.
31. Frank B. Lutz, "The Merits of the Fruit Fly," School Sci. Math., 7 (1907),
672-673.
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thought Payne should try his hand at breeding work, and Drosophila
was handy. "With Morgan I will probably try several things," Payne
wrote Charles Zeleny, "for I fear some will give no results. One
thing I expect to do is to try to rear some aphids and flies in the
dark. On the flies I expect to try all sorts of things to see whether
I can produce variations. ''35 With mice, such projects would have
been much harder to arrange.
It is striking how many of the people whom Morgan got to
work on Drosophila in the early years were, like the fly itself,
relatively low in academic status. These included Morgan's technicians, Eleth Cattell, Sabra Tice, and Edith Wallace, who were
capable college graduates in biology with distinctly limited career
prospects. They had no choice. The students who worked on
Drosophila tended to be those who had little to lose by investing
in a low-status and uncertain creature: Ann Elizabeth Rawls was
a master's student (M.A. 1912), and Leopold Quackenbush was a
long-term graduate student (M.A. 1901) who was chronically ill
(TB?) and dropped out and died a few years later. 36 (It was
Quackenbush who Morgan hoped might collect snails in Hawaii
while restoring his health through outdoor work.) Morgan even went
so far as to recruit Columbia undergraduates, such as Joseph Lift,
Alfred Sturtevant, and Calvin Bridges - his bottle washer.
Ambitious graduate students chose less-risky and higher-status
projects that were more likely to lead to publications and careers.
Fernandus Payne, for example, chose to do his dissertation with
E. B. Wilson on spermatogenesis in toad-bugs. It was sure to pay
off; Drosophila was not.
Quite a number of Morgan's early drosophilists were college
teachers who did their thesis research on the job, over a period
of years. Gail Carver and John Dexter, for example, were professors of biology at Mercer College in Macon, Georgia, and Northland
College in Ashland, Wisconsin, when Morgan recruited them to
work on Drosophila; Roscoe Hyde was professor of biology at
Indiana State NormalY Clara Lynch did her Drosophila work as an
instructor at Smith College, and Shelley Safir did his while
teaching biology in New York City high schools. Mark Stark began
35. F. Payne to C. Zeleny, October 7, 1907, CZ 2. Payne did similar experiments with Wilson, irradiating tadpoles and other organisms in the hope of making
chromosomes segregate in abnormal ways.
36. Catalogue of Officers and Graduates of Columbia University, 16th ed.
(New York: Columbia University, 1916).
37. John Dexter, "On Coupling of Certain Sex-Linked Characters in
Drosophila," Biol. Bull., 23 (1912), 183; Roscoe Hyde, "Fertility and Sterility in
Drosophila ampelophila," J. Exp. Zool., 17 (1914), 141-171.
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Drosophila: A
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it.' ,,49 But just a few days later, Morgan observed a few flies with
a darker pattern than any he had seen before. Inbreeding quickly
produced a mutant strain, with, which had a distribution of
pigmentation that was distinctly higher than the wild type, and
which further selection did not alter. It was just what Morgan had
expected the process of speciation by mutation to look like! Visiting
his wife and newborn daughter in the hospital in the second week
of January, he could talk of nothing but his new mutant. No wonder:
he thought he was witnessing the beginning of a "mutating period"
- evolution induced by strenuous selection in the laboratory, where
all could witness it! 5~ A second mutant, super-with, appeared in
November 1910, thirty generations later.
Morgan's various efforts to induce mutation by altering the
physical environment and by selection were preparations, I believe,
for a major project in experimental evolution, parallel to regeneration, sex determination, and Mendelizing. The pattern of behavior
is unmistakable. He wrote his 1903 book and subsequent articles
in 1905 and 1909 to assimilate current work on adaptation and evolution and to stake out a personal position on key issues. He did
experiments on insects and Drosophila to get hands-on experience of the ways that experimental evidence about variation was
produced. This was just how he had broken into the neo-Mendelian
game, criticizing Cutnot and getting a feel for mouse breeding.
By 1909 he had a distinctive view of variation that served as a
practical guide for breeding experiments, and he thought that
Drosophila might be the ideal organism for experiments on
evolution, just as mice were for Mendelizing and Phylloxera was
for sex determination.
Morgan chose Drosophila for experimental evolution, rather than
some domesticated plant or animal, precisely because Drosophila
was not domesticated. He had long believed that experiments on
evolution would have to be done with wild creatures. "Until we
know more about the results when wild varieties are crossed with
their wild species or with other varieties," he wrote in 1909, "we
can not safely apply Mendel's laws to the process of evolution.''51
49. R.G. Harrison, "Embryology and Its Relations," Science, 85 (1937), 370.
50. Bridges and Morgan, Third-Chromosome Group (above, n. 41), pp. 31-37;
Allen, Morgan (above, n. 8), pp. 148-153. Morgan's daughter was born on January
5, 1910. Lillian Morgan recalled her husband talking of the white mutant, but it
must have been with." T. H. Morgan, "Hybridization in a Mutating Period in
Drosophila," Proc. Soc. Exp. Biol. Med, 7 (1910), 160-161; Elof Axel Carlson,
"The Drosophila Group: The Transition from the Mendelian Unit to the Individual
Gene," J. Hist. Biol., 7 (1974), 37-38.
51. T.H. Morgan, "Breeding Experiments with Rats," Amer. Nat., 43 (1909),
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It is also clear why Morgan did not use Lutz's domesticated stocks
of Drosophila but instead got fresh material from the wild: if he
believed that Lutz's inbred stocks had already entered into a
"mutating period," he would have wanted to avoid such stocks in
experiments designed to see if artificial selection could induce such
an event. (Morgan also insisted that Payne use a wild stock for
his selection experiments, not one from Lutz, doubtless for the same
reason.) 52 Drosophila's natural wildness and lack of Mendelizing
characters, which so limited its use for genetic experiments, made
it the organism of choice for Morgan's new program in experimental
evolution. Morgan meant Drosophila to colonize a new niche in the
ecology of his lab.
DROSOPHILA TAKES OVER
The appearance of with in January 1910 was just the harbinger,
as it turned out, of a gathering flood of mutants: olive body color
and speck wing axil in March, beaded and another olive in May,
along with the famous white eye mutant. Rudimentary wing
appeared in June, pink eye in July, miniature and truncate wings
in August, and at least six more by the end of the year. It appeared
to Morgan that his selection experiment had indeed inaugurated a
de Vriesian "mutating period" in his flies. The character of these
mutants, however, was not quite what the theory predicted. Only
the olive mutants were like with, definite but within the range of
normal variation. The others were more extreme, more like de
Vries's Oenothera mutations or Mendelizing traits. Although they
did not display Mendelian ratios, these mutants did segregate. And
white eye, to Morgan's astonishment, was sex-linked (that is, it
appeared only in males, which had therefore to have a single sex
chromosome). This freshet of extreme mutations was not what
Morgan had expected of a "mutating period." Indeed, it entirely
upset his accustomed way of thinking about evolution and heredity.
Drosophila completely overturned the hierarchy of experimental
lines and organisms in Morgan's laboratory. With the appearance
of white and other Mendelizing mutants, Drosophila broke out of
its restricted domain of experimental evolution and began to take
over Morgan's other experimental lines. The sex-lined character
of white made it an ideal organism for work on sex determina183; Morgan, Evolution and Adaptation (above, n. 43), p. 278. Morgan crossed
wild black and Alexandrian rats in 1908-09.
52. Bridges and Morgan, Third-Chromosome Group (above, n. 41), p. 225;
F. Payne to A. H. Sturtevant, October 16, 1947, AHS 3/19.
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tion, better even than Phylloxera, which was soon set aside for
studies of sex chromosomes and sex determination. Similarly, the
fact that white and other mutants displayed a more or less
Mendelian inheritance made Drosophila the organism of choice
for experimental heredity, better than mice or rats with their limited
number of characters. Mice were soon displaced from their favored
niche in the Columbia laboratory. In short, the proliferation of
mutations in Drosophila altered the domestic ecology of experimental organisms and disciplines. Drosophila began to take over
Morgan's entire operation, displacing other standard organisms in
his other lines of work and carrying with it the new genetic concepts
of genes and chromosomes and the new experimental mode of
genetic mapping. 53
Experimental evolution, ironically, was the one line that did
not survive Drosophila's take-over of Morgan's laboratory. As it
gradually became clear that the flood of mutants resulted simply
from scaled-up production and not from a de Vriesian "mutating
period," selection experiments ceased to have any relevance to
the mechanism of evolution. Besides, Morgan was not one to miss
an experimental windfall, and the Mendelian analysis of mutants
like white was the kind of windfall that happens seldom in a
lifetime. He did continue selection experiments on with and other
indefinite mutants for a year or so, but his gradual abandonment
of these experiments for genetic mapping and linkage marks his
shift from experimental evolution to genetics. 54
Thus, the invasion of Drosophila mutants turned Morgan's
orderly hierarchy of experimental organisms and disciplines topsyturvy. Drosophila just took over its new ecosystem, displacing
53. T. H. Morgan: "Chromosomes and Heredity," Amer. Nat., 44 (1910),
449-496; "Sex-Limited Inheritance in Drosophila," Science, 32 (1910), 120-122;
"The Application of the Conception of Pure Lines to Sex-Limited Inheritance
and to Sexual Dimorphism," Amer. Nat., 45 (1911), 674-678 (this talk was delivered in December 1910); Alien, Morgan (above, n. 8), pp. 137, 148-152.
54. Morgan's last brief fling with experimental evolution began in November
1912 with the appearance of streak, a slight and highly variable thorax pattern
mutant. Bridges and Sturtevant would have discarded it as useless for mapping,
but since it had turned up in a stock that had never been subjected to selection,
Morgan saw a chance to prove what everyone now believed, that selection did
not induce mutations of the with type. He worked on streak for six months, "not
very vigorously" and to little effect. The next such mutant that appeared, trefoil
(November 1913), was discarded forthwith, and in June 1914 all the with stocks
were thrown out. Bridges and Morgan, Third-Chromosome Group (above, n. 41),
pp. 31, 36-44; Calvin B. Bridges and T. H. Morgan, "The Second-Chromosome
Group of Mutant Characters," in Contributions to the Genetics of Drosophila
Melanogaster (Washington, D.C.: Carnegie Institution of Washington, publication no. 278, 1919), pp. 136-140, 222, 244-245.
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established organisms and imposing a new order on the organization of experimental work. It changed forever the natural history
of experimental biological laboratories.
DROSOPHILA: A BREEDER REACTOR
The fundamental cause of this change in experimental practice
was the appearance of large numbers of mutants, and that
phenomenon was in turn caused by the scaling-up of Morgan's
breeding experiments. The more flies that were bred and inspected,
the greater was the probability that mutants would turn up. At some
threshold of scale (five or ten thousand, maybe?) a selection
experiment would have contained enough spontaneous mutants
for one to be noticed. Doubtless many mutants were discarded
unnoticed, but once the first was observed others were more likely
to be picked up - hence the apparent suddenness of their appearanceY But if it was just a phenomenon of scale, why then did Castle
not also observe mutants, since he was working on a scale at least
as large as Morgan and Lutz? The answer is simple: Castle was
working on fertility, and to reduce the burden of routine he counted
only pupae. As a result he worked with only a small number of
adult flies and so did not have a chance to spot mutants! 56
Mutants could have turned up only after Morgan and Lutz began
to work on experimental evolution, since only these experiments
were big enough to cross the statistical threshold. Scaling-up was
crucial. Morgan's early efforts to induce mutation by heat, cold,
or X rays were doubtless done with cultures of a few hundred
flies, and that would not have been enough. It was only in the
inbreeding experiments with thorax pattern, which involved many
generations of mass cultures, that mutants were bound sooner or
later to be spotted. Mendelizing experiments might have been big
enough to cross the statistical threshold, but there was no reason
to use Drosophila for such work, as we have seen. That is why
Drosophila first found a habitable and expandable ecological niche
in experiments on evolution. It was only in this line that the
potential of Drosophila for mutation could be revealed.
Once the threshold of scaling-up was crossed, the production
55. It may also be that the crossing of inbred and wild stocks increased the
natural rate of mutation through an exchange of extrachromosomal genetic elements
or transposons. I am indebted for this idea to Professor Edward B. Lewis. See James
F. Crow and William F. Dove, "The Genesis of Dysgenesis," Genetics, 120 (1988),
315-318.
56. Castle, "Memorandum on the Beginning of Genetic Studies" (above, n.
7).
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ROBERT E. KOHLER
with vermilion, cut, and notch. The Morgan group began speaking
of a "mutating period," perhaps by then as a joke on themselves
these epidemics were not de Vriesian "mutating periods," of
course, but a property of the new mode of practice.
The epidemics were created in part by the drosophilists themselves. Finding a new mutant opened the drosophilists' eyes to
others like it. They began to perceive as mutants, appearances that
they would once have passed over as random variation. Having
created a class of jaunty wings, for example, "suddenly in certain
other stocks wings that have been passed over as 'imperfectly
unfolded' or only vaguely recognized as 'queer' are seen to be
sharply characterized 'jaunties. '''61 "Epidemics" were also in part
the result of imperfect laboratory technique and a certain frenzy
of enthusiasm that afflicted mutant hunters. Many of the mutants
in these "epidemics" turned out to be duplicates, resulting from
accidental contamination of breeding stocks, or simply extreme
forms of natural variation in the wild type - phantoms.
Nevertheless, genuine new mutants kept appearing at an accelerating rate: at least ten in 1911, and two dozen in 1912. By 1915
so many mutant stocks were being cultivated that many of the
less-useful ones had to be discarded; there were just too many to
keep. Such was the profligate fecundity of Drosophila in its new
domestic ecosystem.
Morgan and his drosophilists did indeed discover a "mutating
period," but it had nothing to do with evolution in nature - it was
an aspect of the natural history of experimental laboratories and the
evolution of experimental practices. The awesome productivity of
Drosophila was not simply a physiological property of the organism
itself, though it was partly that. Rather, it was a property of a system
of experimental mass production and of the symbiotic relationship between flies and fly people. The capacity to produce mutants
was the property of a creature with a fast life cycle and large
families when it was brought indoors into a laboratory and integrated into a system for doing large-scale breeding experiments.
The breeder reactor was the whole system of production: the constructed mutant stocks and standard recipes, drosophilists' research
agendas and ways of life. This symbiotic relationship transformed
Drosophila physically into a new domesticated creature, one that
did not exist in nature and that could only have been created in
the peculiar ecology of a genetics laboratory. And so, too, did it
transform the fly people into a new variety of experimental biol-
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More specialized kinds of Drosophila, like fungus- or pollenfeeders, proved almost impossible to cultivate in the laboratory
because they depended on a special local habitat. Similarly, some
wild tropical species, which shunned mankind, were in captivity
highly susceptible to epidemics. Melanogaster was just ideal for life
in the laboratory.
Was it merely an accident that melanogaster was both the first
Drosophila to enter the laboratory and the best suited for laboratory life? Perhaps not entirely. Melanogaster had, after all, been
cohabiting with and adapting to humankind for some thousands
of years. Perhaps there was less difference than we might imagine
between the ecology of the laboratory and of the places, outdoors
and indoors, where melanogaster had long flourished as a hangeron of humankind. The very traits that made melanogaster a good
hitchhiker were precisely the same traits that made it a good
standard laboratory animal. It was hardy and not fastidious about
what or where it fed, and was thus more likely to turn up in
drosophilists' traps and to flourish in overcrowded laboratory jars.
It was relatively tolerant of heat and cold from following humankind
into inclement regions and wintering over in chilly fruit stores
and cellars. It was already an indoor creature, accustomed to human
habitations and able to brave hostile urban environments. It was the
ultimate successful opportunist. Cosmopolitan species, in short,
were preadapted to laboratory life, and D. melanogaster was the
cosmopolitan fly par excellence. It had been shaped by natural
selection to live in a commensal relationship with humankind. It
was the most likely to turn up on a window ledge or in a trap.
The forces that made melanogaster such a successful cosmopolitan
and camp follower of the most cosmopolitan of primates, also suited
it well to a symbiotic relationship with the variety of humankind
who inhabit experimental laboratories.