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Mendel and modern genetics:

the legacy for today
Garland E. Allen
Department of Biology, Washington University, St Louis, MO 63130, USA

The legacy of Mendels pioneering studies of hybridization in the pea continues to influence the way we
understand modern genetics. But what sort of picture
did Mendel himself have of his work and its ultimate
uses, and how does that picture compare with the collection of ideas and methodologies that was put forward in his name and later became known as
Mendelism? With genetics standing at the center of
our present biomedical and biotechnological research,
an examination of the history of our concepts in the
field can help us better understand what we should and
should not expect from current genetic claims. For that
enterprise there is no better starting place than Mendel
himself.
As we celebrate 50 years of the Watson Crick model of
DNA, it is worth stepping back to take a longer look at the
work that started it all: the pioneering hybridization
studies of the pea (Pisum sativum) and bean (Phaesolus)
by Gregor Johann Mendel (1822 1884) (Fig. 1). Although
Mendels work was first presented at a meeting of the
Brunn Natural History Society in 1865, and published in
its Proceedings in 1866, it received only a modicum of
attention until 1900, when it was more or less simultaneously rediscovered by three different investigators:
Carl Correns (1864 1933) in Germany, Hugo De Vries
(1848 1935) in the Netherlands, and Erich von Tschermak-Szeneygg (1871 1962) in Austria. Whilst all three
found Mendels work suggestive, it is not clear that any of
them really saw the significance of what he had done, and
none of the rediscoverers became a major promoter of the
new genetics [1].
The first major publicist for Mendels work was William
Bateson (1865 1926) in England. Through the Royal
Horticultural Society, Bateson had Mendels paper translated into English for the first time, and wrote a general
exposition that laid out the basic principles of what soon
came to be known as Mendelism [2]. Batesons work
brought Mendel to the attention of numerous workers in
England, Scandinavia and the United States, in particular
to many of those involved in practical animal and plant
breeding [3]. Although there was reluctance in some
quarters to embrace Mendels work immediately
especially among academic biologists by the end of the
first decade of the 20th century, the theory had gained a
considerable following. However, the explanation Mendel
Corresponding author: Garland E. Allen (allen@biology2.wustl.edu).

offered for his breeding data seemed reminiscent of so

many of the speculative, particulate theories of heredity
that had abounded in the post-Darwinian era, including
Darwins own provisional hypothesis of pangenesis,
August Weismannns elaborate theory of ids, idants and
biophors, Ernst Haeckels imaginary plastidules, and
Hugo de Vries postulated pangenes. Consequently, to
some, at least, Mendels work seemed like just one more of
the sort of abstract proposals they had encountered all too
frequently. In point of fact the paper probably seemed
extraordinarily dense, with no illustrations but numerous
binomial expansions, which were likely to have been a putoff for many biologists who at the time were notoriously
math-shy. For whatever reason, for the first decade of its
re-emergence into the scientific world, Mendels contribution remained controversial at best, dismissed by
significant sections of the biological community at worst.
What ultimately served to establish Mendelism on more
firm ground between 1900 and 1915 was: (1) its extension
to an increasingly wide variety of organisms; and (2) its
unification with the cytological work on chromosomes
carried out principally through the work of Thomas Hunt
Morgan (1866 1945) and his young, enthusiastic team of
investigators at Columbia University between 1911 and
1925. The work of the Morgan school demonstrated that
the abstract elements or factors discussed by early 20thcentury Mendelians could be regarded as discrete,
material units arranged linearly along the chromosomes,

Fig. 1. Gregor Johann Mendel (standing, second from right) with members of the
Augustinian monastery of StThomas in Brno in about 1862. Mendel can be seen
observing a plant specimen. Others in the photograph of particular importance in
Mendels life are the Abbot Cyrill Napp (seated, second from right) and Matous
Klacel (seated, first from right), who was also interested in natural science and
philosophy, and with whom Mendel had frequent lively discussions. Reproduced,
with permission, from [5], p. 212.

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Box 1. The common picture of Mendelian theory

Mendel observed the inheritance patterns of traits or characteristics in pea plants, such as height, pod color and or seed shape,
each of which showed alternate forms: tall/short, yellow/green
and smooth/wrinkled, respectively.
Mendel referred to these alternate conditions as dominant and
recessive.
Mendel hypothesized that each trait was represented in the germ
cells of adult plants by two determinants (referred to in his paper
as Anlagen or elements), one received from each parent; these
determinants were symbolized by Mendel with a capital letter for
the dominant form (e.g. A) and a lower-case letter for the recessive
form (e.g. a).
The determinants could be combined in one of three ways: two
dominants (AA), two recessives (aa) or a dominant and a
recessive, or hybrid (Aa).
Although he knew nothing about the cytology of chromosomes,
Mendel hypothesized that in the formation of the pollen or egg
cell, the two factors for each trait would separate and go into
different gametes; thus a parent that was pure dominant would
produce gametes all of which contained the dominant factor (A),
and a parent that was pure recessive would produce gametes all of
which contained the recessive factor (a); hybrid parents, however,
would produce two kinds of gametes: 50% would contain the
dominant factor (A) and 50% the recessive factor (a).
At fertilization, the double-determinant condition would be
restored.
If both parents were hybrids, any of the three possible combinations could occur and would be distributed randomly, according to the laws of probability: 1 AA: 2 Aa: 1 aa; because organisms
that are Aa and AA look alike, the ratio based on appearance of the
traits (what later came to be called phenotype) would be 3:1.
When two or more characteristics (e.g. Aa, Tt) are followed in a
dihybrid cross, the two sets of determinants segregate randomly,
so that any combination of A, a, T and t is possible, what came to be
known as the principle of random assortment.
Factors somehow determine traits, so that after the term gene
was introduced in 1909, it was common to speak of a gene for
tallness or a gene for wrinkled seed. During the early years of the
Mendelian theory, this was referred to as the unitcharacter
hypothesis.
Factors are not modified by being combined with their alternate
form; thus, the factor, t, for shortness is not affected in any way by
being combined with the dominant factor, T, for tallness in the
hybrid, a concept that became known as the concept of the purity
of the gametes.

and that observed variations in the patterns of inheritance

of traits could be traced to the mechanics of chromosome
behavior during meiosis (gamete formation). Mendels
factors (after 1909 referred to as genes) thus seemed to be
real, material units, not metaphysical postulates. In
recognition of this work, Morgan was awarded the first
ever Nobel Prize for research in genetics in 1933 [4].
Therefore, a common picture of Mendelian theory has
emerged from this early work and has been promoted in
textbooks ever since (Box 1).
This scheme has made for a very heuristic pedagogy,
and has been replicated in an almost infinite number of
high-school and college biology textbooks. Although it has
enjoyed a certain intellectual neatness, this formulation
has several problems that have now begun to surface as
molecular genetics has provided a far more sophisticated
understanding of gene function than was available in the
first half of the 20th century. The first problem is simply
historical: it is not clear that the neat textbook scheme is
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the sort of picture Mendel himself envisaged. Second, as an

introduction to the study of genetics, it has led to
unfortunate and now long-entrenched misunderstandings
of how genes really function, and the relationship between
genes and the development of adult traits that has carried
over into molecular genetics.
Mendels background
Mendel was born on 22 July 1822, in the small rural village
of Hyncice, in Moravian Silesia, then part of the Austro
Hungarian Empire. As the only son of a peasant farmer, he
was expected to follow in his fathers footsteps and take up
farming. But early on, his interest in natural history and
his studious ways brought him to the attention of the
priest, Friar Schreiber and the local schoolteacher. In spite
of financial hardship for his family, young Johann was sent
to a larger school in a nearby village and eventually
qualified for Gymnasium in Opava (Troppau). It was a
period of extreme privation, but Mendel managed to
graduate in 1840 with considerable academic success. One
of the chief influences on him at the time were Schreibers
Enlightenment ideals, particularly his emphasis on
science as a way of dispelling superstition and ignorance.
Schreiber was keenly interested in applying scientific
principles to improve humanity, and was heavily involved
with local agricultural groups and the Pomological
Society [5].
Financial problems continued to plague Mendel as he
tried to continue his studies at the Philosophy Institute in
Olomuc (Olmutz). After several periods of illness, brought
on it is thought by his poverty and overwork, Mendel
managed to finish his studies at the Institute and entered
Olomuc University. Here, according to records, he took a
course of lectures in physics, mathematics and logic.
However, unable to complete his degree owing to financial
constraints, he applied for, and was accepted into, the
Augustinian monastery of St Thomas at Brno (then
Brunn), Moravia in 1843 (Fig. 2). Although he did not
feel a particularly fervent spiritual calling, Mendel
realized shortly after joining the monastery that at last
he was free from constant financial worries and could
pursue his intellectual interests in exchange for attending
to pastoral duties.
The monastery at Brno was a center of learning in the
early- and mid-19th century, especially in the natural
sciences and agriculture [6]. The practical and economic
benefits of promoting new agricultural practices was
among the monasterys top priorities, with most of its
income coming from extensive landholdings leased to local
farmers. At the time that Mendel entered St Thomas,
the Abbott (administrator) was Friar F. Cyril Napp
(1792 1867), an enthusiastic naturalist, member of
several local agricultural and scientific societies, and
author of many technical papers, particularly on plant
pests. In 1830, he had given over one part of the monastery
garden to another monk, Matous Klacel (1808 1882) for
experimental cultivation of rare Moravian plants. The
monastery also had an extensive library containing a
variety of scholarly texts (Fig. 2).
The importance of agricultural concerns in Mendels
developing interests in the natural sciences is clear [7]. At

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Fig. 2. The main church and monastery building (left) that became the seat of the Augustinian Order in 1783, and the magnificent library of the monastery (right), subsequently the Mendelianum, a division of the Moravian Museum. One of Mendels early mentors in the monastery was Matthew Kla cel, who became, after demotion from
position of teacher because of his radical ideas, the monasterys librarian. The library has many works on agriculture, including Liebigs 1844 Agricultural Chemistry, and
Mendels copy of the 1860 German edition of Darwins On the Origin of Species. Reproduced, with permission, from [5], p. 46.

the time, the amount of experimental breeding in Moravia,

both with plants and animals, was extensive. Indeed, the
Brunn Natural History Society, before which Mendel read
his Pisum paper in 1865, was a section of the larger Brunn
Agricultural Society. The peas with which Mendel worked
were derived from major edible strains, suggesting the
close connection between his studies on hybridization and
agricultural interests. Mendel clearly benefited from the
monasterys support of science, and the emphasis placed
on agriculture by local agriculturalists and members of the
monastery staff turned Mendels interests in a specific
direction that ultimately led to his extensive series of
breeding experiments.
Motivation for Mendels Pisum experiments
It was during his period as a teacher at the newly opened
Realschule that Mendel began his systematic breeding
experiments with Pisum. One of the major reasons that
has been put forward to explain why a clergyman and
schoolteacher would have undertaken such an unusual
investigation is that he wanted to provide a generalized
theory of heredity that would complement Darwins theory
of natural selection (or, as several authors have also
suggested, to counter Darwins theory by demonstrating
the fixity of species limits during hybridization). Darwins
theory had encountered considerable theoretical difficulties because of the prevailing belief by naturalists and
breeders (including Darwin himself) in blending inheritance the idea that any trait in the offspring was a
blend of the traits observed in the two parents. Thus, if
parental differences blended in the offspring, new variations would be diluted in every generation, leaving little
for natural selection to act on. However, it is clear that
Darwins work could not have been Mendels initial
motivation, because he had already begun his experiments
in 1856, three years before the publication of The
Origin of Species.
If not evolution, what was the initial motivation for
Mendel, and what did his paper say, both to his
contemporaries and to those who read him after the
rediscovery in 1900? In 1979, Robert Olby revised our
understanding of the Pisum paper by stripping it of its
20th-century interpretations [8]. He concluded that
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Mendel was not a Mendelian in any modern senses of

term, and that proponents of Mendelism, from Bateson
onward, had read much into Mendels work that was not
part of his original formulation. Similar re-evaluations
have subsequently been offered by others [9 11].
The gist of these various reinterpretations is that:
(1) the primary motivation for Mendels work was not to
develop a generalized theory of heredity, but was far more
immediate and practical to establish patterns of
hybridization that would have been of interest to agricultural breeders and others in 19th-century Moravia; and
(2) Mendel never proposed that there were material
particles, factors or Anlagen, in the germ cells, or that
such particles were necessarily transmitted via pollen and
egg cells to the offspring. Mendel, they argue, remained
far more agnostic about the physical basis of inheritance
than later Mendelians assumed.
Indeed, it is much more likely that the direction of
Mendels hybridization research was guided by the strong
agricultural interests around him. It was precisely
through hybridization that breeders hoped to generate
new combinations of characters that they could exploit to
form new breeds. The problem was that most hybrids do
not breed true and have a tendency to revert to their
original parental types. What Mendel was able to explain
were the conditions under which that happened, and most
importantly, the methods for distinguishing between
hybrids and true-breeding forms that might look phenotypically similar. Crucially, his hypothesis of dominant and
recessive characters provided an alternative to blending
inheritance. Whatever happened within the germ cells of
the hybrids, the characters were not blended by residing
together. This became as important a principle for breeders
as it would become later for Darwinian evolutionists,
because it freed both groups from the loss of distinct
variations through blending or swamping in the hybrid.
Nevertheless, it is not possible to be interested in
hybridization without simultaneously being drawn into
accepting, however loosely, some concept of heredity.
Hybridization involves mixing germ lines from two
different ancestries, and how those germ lines interact,
by definition, forms a theory of heredity. Thus, to claim
that Mendel was interested only in hybridization and not

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in any general principles of heredity is to make a confusing

distinction. The two processes are not only not mutually
exclusive, they are complementary. The revisionist view
therefore is that Mendels major motivation (and interest)
lay in establishing principles of hybridization, and not in
establishing a generalized theory of heredity as was
assumed by his early-20th century followers.
These re-interpretations of Mendel also argue that
nowhere does he explicitly state that the germ cells contain
particles that determine the character of a trait. Mendels
symbols (e.g. T,t and A,a), they argue, were merely
algebraic notations and were never meant to represent
material entities in any biological sense. It is true that
Mendel is not explicit on this matter. He talks in his paper
about alternate characters (tall and short) segregating in
the germ cell of the hybrid parent, but he does not refer to
determiners or particles; moreover, he does not talk about
segregation of the two characters in the homozygous
forms, and represents homozygotes symbolically with only
a single letter (such as A for the dominant parent or a for
the recessive), whereas he always represents the hybrids
with two letters (e.g. Aa). Thus, he consistently shows the
results of a monohybrid cross as A 2Aa a.
A careful re-reading of the Pisum paper suggests that
Mendel was ambiguous, but not explicitly agnostic,
towards the notion of hereditary particles. The ambiguity
arises because throughout his paper, Mendel consistently
talks about the characters [Charactere ] of the parents or
offspring of his crosses, which of course refer to the visible
adult traits. His empiricism and training in the physical
sciences led him to emphasize what he could see in the
plants he was breeding. At the same time, toward the end
of his paper, Mendel wrote about the implications of his
experiments for understanding the composition of the
fertilization cells (i.e. pollen and egg). Here, he introduces
the terms Anlage and Elemente in reference to what is
transmitted from parent to offspring through fertilization.
The development proceeds from a constant law which is
grounded in the material composition and arrangement of
the elements [Elemente ], which come together in the cell in
a viable union, wrote Mendel [12].
Indeed, it would have been unusual in Mendels day,
and given his training, not to at least have thought in
terms of some sort of discrete hereditary particles being
passed from parent to offspring during reproduction.
Virtually every theory of heredity proposed during the
middle and later years of the 19th century was couched in
particulate terms so that it was common among biologists,
especially in the German-speaking world, to think in
terms of atom-like particles or molecules as agents of
hereditary transmission. The problem with most of these
theories was that they were largely speculative, and based
on a few observations and little, if any, experimental
evidence. It is not surprising therefore, that Mendel hints
at the existence of such components in the germ cells.
However, it is important to remember that he was first and
foremost an empiricist, and was highly restrained when it
came to speculation about mechanisms. In this sense,
historians are right who emphasize that Mendel was
primarily interested in establishing the laws of hybridization rather than a general theory of heredity. His
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constant emphasis on characters, his recognition that it

was sometimes difficult to determine in what character
class an individual should be placed, and his meticulous
record-keeping all testify to his strongly empirical and nononsense approach to experimentation.
Mendelism and the unit character hypothesis
By the time Mendels paper was rediscovered in 1900 and
had begun to receive attention from biologists and
breeders in industrialized countries, economic, social and
intellectual conditions were significantly different than
those in 1866 Central Europe. The industrial revolution in
Europe and the US had placed many new demands on
agriculture [13]. Not only were there increasing demands
on food production to supply the large urban workforces
that had developed in industrial areas, but much of that
workforce had come from the agricultural sector, leaving it
in short labor-supply. Mechanization of agriculture had
proceeded with great rapidity at the end of the 19th
century, a process that had initiated what has been called
the industrialization of agriculture [14]. During the
second half of the 19th century, scientific agriculture
based on the organic and physiological chemistry of Justus
von Liebig (1803 1873) and his school in Giessen had
greatly improved yields by attention to aspects such as
fertilizers and animal and plant nutrition. But by 1900,
those inputs had, for the time, achieved their technical
limits. Improvements in breeding higher-yielding varieties,
however, held out a wholly new potential.
One aspect of agriculture husbandry was still in a
rudimentary stage at the turn of the 20th century. Most
animal and plant breeders operated by various rules of
thumb that they had developed their separate localities or
had learned and modified from others. It was largely a
craft, with no general rules that applied across the board.
The reintroduction of Mendels work at this juncture
generated hope that some principles of heredity might, at
last, provide the breeder with methods that could yield
more predictable results. Although in reality the application of Mendelian principles to improving animal and
plant productivity turned out to be more difficult than it
initially seemed, optimism was nonetheless shown by
many academic biologists, US Department of Agriculture
(USDA) officials, and breeders in the early decades of the
century. The economic and social conditions made Mendels work look profitable in 1910 in a way it had not looked
in 1866.
But as academics and breeders took up the nascent
science of Mendelism [15], there remained an uncertainty
over what was actually transmitted from parent to
offspring during fertilization. Although Wilhelm Johannsen (1857 1927), the Swedish plant breeder who coined
the term gene, and William Bateson, who coined the term
genetics, both favored a more abstract statistical model of
the hereditary unit, others, especially in the US, pressed
for a more material, atomistic interpretation from 1903
onward. The abstract element of Mendels paper had
quickly become the discrete gene of the rediscovered
Mendelism. Textbooks began to present images of
Mendelian crosses using Mendels capital and small letters
for dominant and recessive traits.

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Between 1900 and 1910, this interpretation of Mendelism communicated two important notions to the generation of biologists who were beginning to learn about the
new genetics. First, there was no hard-and-fast distinction
between the letter or factor and the adult character for
which it stood. Thus, geneticists would write about the
inheritance of height or wing shape or red eyes as if the
trait itself was what was carried in the male or female
gamete. Furthermore, in the early decades after the
rediscovery of Mendelian theory, the identification of the
hereditary unit with the adult trait was graphically
reinforced by the Punnet Square a method of calculating
Mendelian combinations which blurred what would
become known after 1911 as the phenotype genotype
distinction, and gave the impression that the inherited
factor was the trait in miniature. It was this conflation that
led embryologists like the young T.H. Morgan to reject the
Mendelian hypothesis for a decade [4]. For Morgan and
others, the Mendelian factor or gene smacked too much
of the embryologists old bugbear, preformationism (the
idea that the complete adult exists already preformed in
the fertilized egg, and that embryonic development
involves only an unfolding or growth of the embryo in
size). Epigenesis, the alternative view, had replaced
preformationism by the mid-19th century, so that Mendels
work in the form presented by Mendelians, seemed like an
outdated throwback to a long-discarded idea.
With the wedding of Mendelian theory to the cytological
investigation of chromosomes by Morgan and his group
after 1910, the material, discrete and atomistic concept of
the gene gained considerable prominence. By carefully
correlating the inheritance patterns of gene mutations
with observable changes in chromosome structure, Morgan and his students were able to show that genes could be
clearly regarded as material entities that occupied specific
positions loci linearly arranged along the chromosome.
By 1915, when the group published the their pathbreaking book, The Mechanism of Mendelian Inheritance
[16] the beads on a string model of the chromosome had
become an icon. Not only did this model promote the idea of
genes as atomistic units, it also further supported,
indirectly, the unit character hypothesis. The gene was
now viewed not only metaphorically, but also literally as an
atom, entering now into this, now into that combination,
but emerging each time with its own integrity in the same
fashion as atoms entered into, and came out of, molecular
combinations. Indeed, the language of chemistry began to
enter genetics explicitly in relation to the discreteness of
the Mendelian factor. Harvard geneticist W.E. Castle
(1867 1962) wrote early on that:
all observed inheritance phenomena can be expressed satisfactorily in terms of genes, which are supposed to be to heredity what
atoms are to chemistry, the ultimate, indivisible units, which
constitute gametes much as atoms in combination constitute
compounds [17].

Whilst Castle was arguing that genes were not quite like
atoms because they could be altered by the selection
process, that he mentions the analogy at all indicates
something of its prevalence. Somewhat earlier his colleague E.M. East had written that Mendelism is therefore
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just such a conceptual notation as is used in algebra or in

chemistry [18], whilst British polymath J.B.S. Haldane
(1892 1964), as late as the 1930s, claimed that the
atomic nature of Mendelian inheritance suggests very
strongly that even where variation is apparently continuous
this appearance is deceptive. On any chemical theory of
the nature of genes this must be so. [19]
The implication that the discreteness of the gene
implied the organism was constructed as a mosaic of
adult traits was given explicit voice by Bateson within the
first years of his encounter with Mendelism. In 1901, he
wrote:
In so far as Mendels law applies, the conclusion is forced upon us
that the living organism is a complex of characters of which some,
at least, are dissociable and are capable of being replaced by
others. We thus reach the conception of unit characters, which may
be rearranged in the formation of reproductive cells [20].

The next year Bateson wrote even more boldly: The

organism is a collection of traits. We can pull out
yellowness and plug in greenness, pull out tallness and
plug in dwarfness. [21] This mosaic view of the adult
organism parallels the mosaic view of the genome, and
thus pictures the organism as an aggregation of interchangeable parts that can be arranged and rearranged by
assembling the right combination of genes. Admittedly, not
all early Mendelian geneticists took quite such a mechanical view of heredity, but Bateson was one of the leaders in
the field and among its most vocal proponents. His
influence was therefore considerable. It is thus even
more ironic that as late as 1922, Bateson still held strong
reservations about linking Mendelian genes to chromosomes [22].
The atomistic, mosaic conceptualization of the gene was
initially fruitful, allowing biologists to develop a quantitative, experimental and predictable aspect of their science
that made it as hard as anything in chemistry or physics
[23]. However, it created problems almost from the outset.
It allowed biologists to put aside questions of gene
function, or the relationship between Mendelian genes,
embryonic development, and even evolution. From 1915
onward, most attention was paid to the mechanics of gene
shuffling during transmission of chromosomes from parent
to offspring. This trend, which marked the hey-day of
classical genetics, led to the extraordinary feat of mapping
the chromosomes of several model organisms, such as the
fruitfly, maize and mouse. But the atomistic trend in
classical genetics also left an unfortunate legacy. By
perpetuating the unit character concept of the gene,
even with lip-service paid to the idea of gene interactions,
the vast majority of the public and many biologists as well,
still hold to the view that one, or at best a very few, genes
determine each specific, adult trait.
Nowhere is the tendency to attribute discrete gene
elements to specific traits more common, and perhaps
more socially troublesome, than in the area known as
human behavior genetics. To date, social and personality
traits as complex and varying as schizophrenia, manic
depression, alcoholism, criminality, violence, shyness,
homosexuality, risk-taking and religiosity, have all been
headlined at one time or another as caused by a gene. Not

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only do such claims greatly oversimplify the biological

process of development, during which genes interact with
other genes and the environment, they suggest a set of
potential solutions that is deceptive: drug therapy, gene
therapy or sterilization. Indeed, sterilization was
implemented in both the US and Germany in the old
eugenics movement, when simplistic genetic notions
allowed the compulsory sterilization laws to be passed
for those with undesirable hereditary conditions [23].
Today, Norplant can replace surgical sterilization, and
pharmacogenetics looms on the horizon, promising to
counteract the effects of defective genes and to yield huge
benefits for the pharmaceutical industry.
Genes are of course critical components in one way or
another of all of our traits. But because of the increasingly
detailed findings of molecular genetics and the Human
Genome Project, the picture of what genes do and how they
function in the development of adult human characters
has changed dramatically from the legacy built out of the
early interpretations of Mendels work. With his skepticism about undue speculation regarding the mechanics of
the hereditary process, Mendel might indeed have been
more in tune with modern genomics than with the
Mendelians who crafted the classical concept of the gene
in his name.

2
3

4
5
6
7

8
9
10

Conclusion
As we look forward, it is clear that one of the most
important forefronts of modern genetic research lies in the
area of what has been variously called the genetics of
development, developmental genetics, and when combined with evolutionary theory, the evolution of development or Evo Devo for short. These emerging fields,
which synthesize the classical genetics of the first half of
the 20th century with the molecular genetics and
evolutionary biology of the second half, promise to yield
a more complex, and hopefully more realistic picture of
how the genome guides the individual organism from
fertilization to adulthood.
We are standing at a divide between an old and a new
genetics, between a mosaic and mechanical and a holistic
and integrated view of the organism that promises to yield
exciting results in the years ahead. It behoves us to
understand the pathway that has brought us to this
historic juncture, not only so that we may take the fullest
advantage of our newest research paths, but also that we
avoid the disastrous social consequences that can arise
from misplaced expectations of what genetics can do. Like
all other scientific and technological findings, we must first
understand the science itself and its history to recognize
both its potential and its limitations.

11

References

23

1 Sturtevant, A.H. (1965) History of Genetics, Harper & Row; in chapter

21, Sturtevant reviews the claims that the three rediscoverers were
really so independent, whilst several papers by Alain Corcos and Floyd
Monaghan suggest that only Correns had really collected data that

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12

13

14

15
16
17
18
19
20
21

22

were really comparable to those obtained by Mendel. See, Corcos, A.

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Tscheremak: a non-discoverer of Mendelism II: a critique. J. Hered. 78,
2 10. These various positions have been admirably summarized in
Peter Bowler (1989) The Mendelian Revolution. The Emergence of
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Bateson, W. (1902) Mendels Principles of Heredity: A Defense,
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Letter from William Bateson to his wife, Beatrice, 3 October 1902,
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Allen, G.E. (1978) Thomas Hunt Morgan, the Man and His Science,
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Olby, R. (1979) Mendel no Mendelian? History of Science 17, pp. 53 72
Brannigan, A. (1979) The reification of Mendel, Social Studies of
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Meijer, O. (1983) The essence of Mendels discovery. In Gregor Mendel
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Monaghan, F.V. and Corcos, A. (1984) On the origins of the Mendelian
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Mendel, G., cited in [11], p. 105. The original reads: Diese Entwicklung
erfolgt nach einem constanten Gesetze, welches in der materiellen
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Allen, G. (2000) The reception of mendelism in the United States,
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On the industrialization of agriculture in 20th-century America, see
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366 367, Holt, Rinehart and Winston; and Kloppenberg, J.R. (1988)
First the Seed. The Political Economy of Plant Biotechnology. Cambridge University Press, especially chapters 2 4
Punnett, R. (1911) Mendelism, Macmillan
Morgan, T.H. et al. (1915) The Mechanism of Mendelian Inheritance,
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Castle, W.E. (1919) Piebald rats and the theory of genes. Proc. Natl.
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East, E.M. (1912) The Mendelian notation as a description of
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Haldane, J.B.S. (1932) The Causes of Evolution, p. 57, Harper and
Brothers
This quotation comes from Punnett, R.C., ed. (1928) Scientific Papers
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Bateson, W. (1902) Mendels Principles of Heredity: A Defense,
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