Mendel: Morphologist and Mathematician Founder of Genetics - To Begin a Celebration of the 2015 Sesquicentennial of Mendel's Presentation in 1865 of his Versuche über Pflanzenhybriden


Opitz, J.M.; Bianchi, D.W.

Molecular Genetics & Genomic Medicine 3(1): 1-7

2015


Entering a school building, recently erected at state expenses, to attend a regularly scheduled evening session of a local and regional scientific association, few would give any thought to its foundation. Except perhaps those who watched the building go up in the downtown area of Brehm (Brno) in 1858/9 and who taught in it, as did Mendel (Fig. 1). Thus, 150 years ago, on a cold clear (IItis 1924) February night (2/8/1865) a small group of Augustinian monks, all priests, Mendel among them, made their way from their more or less permanent abode, the renowned, ancient abbey of St. Thomas, along the Johannesgasse to the new school building to hear Fr. Mendel present the results of his 8 years of research into plant hybrids, specifically crosses of several types of Pisum sativum, garden peas. The monastery with its beautiful Gothic church (Fig. 2) was located in Altbriinn (Stare Brno), incorporated into Briinn, then as now the capital of Moravia. At the time, Moravia was in the Austrian Empire in that portion of Silesia left to Austria after the three wars between Frederic the Great and Empress Maria Theresia. Later it was in Czechoslovakia, and at present is in the Czech Republic.

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Genomic
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INVITED
COMMENTARY
MENDEL:
Morphologist
and
Mathematician
Founder
of
Genetics
-
To
Begin
a
Celebration
of
the
2015
Sesquicentennial
of
Mendel's
Presentation
in
1865
of
his
Versuche
Uber
Pflanzenhybriden
John
M.
Opitz
l
&
Diana
W.
Bianchi
2
'Departments
of
Pediatrics
(Medical
Genetics),
Human
Genetics,
Pathology,
Obstetrics
and
Gynecology,
University
of
Utah
School
of
Medicine,
Salt
Lake
City,
Utah
2
Mother
Infant
Research
Institute,
Tufts
Medical
Center,
Boston,
Massachusetts
doi:
10.1002/mgg3.127
Introduction
Entering
a
school
building,
recently
erected
at
state
expenses,
to
attend
a
regularly
scheduled
evening
session
of
a
local
and
regional
scientific
association,
few
would
give
any
thought
to
its
foundation.
Except
perhaps
those
who
watched
the
building
go
up
in
the
downtown
area
of
Brfinn
(Brno)
in
1858/9
and
who
taught
in
it,
as
did
Mendel
(Fig.
1).
Thus,
150
years
ago,
on
a
cold
clear
(Il-
tis
1924)
February
night
(2/8/1865)
a
small
group
of
Augustinian
monks,
all
priests,
Mendel
among
them,
made
their
way
from
their more
or
less
permanent
abode,
the
renowned,
ancient
abbey
of
St.
Thomas,
along
the
Jo-
hannesgasse
to
the
new
school
building
to
hear
Fr.
Men-
del
present
the
results
of
his
8
years
of
research
into
plant
hybrids,
specifically
crosses
of
several
types
of
Pisum
sati-
vum,
garden
peas.
The
monastery
with
its
beautiful
Gothic
church
(Fig.
2)
was
located
in
Altbrfinn
(Stare
Brno),
incorporated
into
Brfinn,
then
as
now
the
capital
of
Moravia.
At
the
time,
Moravia
was
in
the
Austrian
Empire
in
that
portion
of
Silesia
left
to
Austria
after
the
three
wars
between
Frederic
the
Great
and
Empress
Maria
Theresia.
Later
it
was
in
Czechoslovakia,
and
at
present
is
in
the
Czech
Republic.
Entering
the
Oberrealschule,
the
monks
and
their
secu-
lar
fellow
members
and
guests
of
the
Society
of
Natural
Sciences
(Naturforschender
Verein)
left
their
top
hats,
canes,
and
capes
in
the
lobby,
took
their
seats
close
to
the
stove,
noting
the
officers
of
the
Association
moving
for-
ward
to
where
Mendel
had
seated
himself
in
the
front
row.
The
Vice
President
of
the
Association,
the
distin-
guished
botanist
Carl
(or
Karl)
Theimer,
announced
the
evening's
agenda
and
asked
the
Association's
equally
accomplished
secretary,
the
botanist
and
astronomer
Gustav
V.
Niessl
to
introduce
the
speaker,
Mendel,
and
invited
him
to
give
the
fi
rst
of
his
two
presentations
on
Experiments
in
Plant
Hybridization
(Versuche
hber
Pflanzenhybriden).
The
second,
Mendel's
summary
and
conclusion,
was
given
1
month
later
(3/8/1865).
Thus,
more
or
less
imaginatively
retold,
occurred
an
event,
surely
equal
in
importance
in
the
history
of
Wes
-
I
Figure
1.
Pencil
and
ink
sketch
of
Gregor
Mendel
(1822-1884)
as
imagined
at
the
time
he
gave
his
lecture
on
February
8,
1865.
Used
with
permission
by
the
artist
Claire
Harper
and
provided
by
Dr.
Sherri
Bale,
GeneDx.
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
This
is
an
open
access
article
under
the
terms
of
the
Creative
Commons
Attribution
License,
which
permits
use,
distribution
and
reproduction
in
any
medium,
provided
the
original
work
is
properly
cited.
1
Invited
Commentary
J.
M.
Opitz
&
D.
W.
Bianchi
El
If
1
1
It
II
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i
dlllllllllllllllllllll
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1111111111111
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111
tern
Biology
to
the
presentation
just
7
years
earlier
by
Darwin's
friends
and
colleagues
to
the
Linnean
Society
of
London
(Slotten
2004)
on
the
theory
of
descent
with
modification
through
the
action
of
natural
selection
pro-
posed
by
Wallace
and
Darwin.
Mendel
and
Darwin
In
contrast
to
Darwin,
Mendel,
quite
aware
of
the
theory
of
evolution,
presented
facts,
not
theory,
on
inheritance,
dis-
covered
not
through
natural
selection
but
by
artificial
selec-
tion,
that
is,
by
means
of
plant
hybridization.
When
Darwin
fi
nally
published
in
1859
he
left
a
substantial
tome:
"On
the
Origin
of
Species
by
Means
of
Natural
Selection,
or
the
Preservation
of
Favored
Races
in
the
Struggle
for
Life,"
the
argument
(for
natural
selection)
sustained
by
and
meant
to
persuade
principally
by
the
sheer
weight
of
a
huge
body
of
facts
and
data
and
inferences
drawn
from
them.
The
issue
it
addressed,
was
condensed
into
a
single
word:
"evolution"
urged
upon
a
reluctant
Darwin
by
Herbert
Spencer
and
used
only
once
in
1859
("evolved"),
in
the
last
sentence
of
The
Origin.
Darwin,
a
19th
century
morpholo-
gist,
was
rather
conscious
of
its
use
until
recently
in
an
embryological
context.
Darwin
was
equally
nimble
and
expert
in
adducing
animal
and
plant
examples
in
his
book;
Mendel,
also
a
19th
century
morphologist,
confined
himself
to
plants
in
his
presentation,
although
later
he
performed
experimental
breeding
with
bees.
For
Darwin,
artificial
selection
was
a
means
of
hurrying
up
natural
selection
in
producing
the
immense
variety
of
domesticated
animal
and
plant
forms
from
one
or
more
than
one
ancestor.
Mendel
dealt
with
both,
wild
forms
collected
around
Brfinn
(i.e.,
the
hawkweed,
Hieraciurn,
which
later
caused
him
difficul-
ties
in
validating
the
conclusions
of
his
earlier
experiments)
Figure
2.
Partial
view
of
the
Koniginkloster
("Queen's
Cloister"),
Abbey
of
St.
Thomas
of
Altbriinn
taken
by
Diana
Bianchi
in
2003.
View
from
southwest.
On
the
left,
between
two
trees
is
the
Charlemont
monument
to
Mendel.
The
part
of
the
building
with
clock
tower
houses
the
library
and
Mendel
museum.
The
garden
area
between
fence
and
building
is
the
area
where
Mendel
presumably
had
his
experimental
plot.
Behind
the
clock
tower
is
the
gable
of
the
church
of
St.
Thomas
from
1353.
and
forms
under
cultivation,
mostly
for
practical
agronom-
ical
(decorative
and
edible)
purposes.
Darwin
postulated
that
it
took
millennia
and
hundreds
of
generations
for
nat-
ural
selection
to
lead
to
a
new
species,
while
Mendel
stud-
ied
modification
of
plant
forms
from
well
-established
parent
stock
in
only
a
few
generations.
Mendel
was
fully
aware
of
Darwin's
work,
as
witnessed
by
his
many
annota-
tions
in
the
1863
German
translation
of
Darwin's
works
preserved
to
this
day
in
the
extensive
monastery
library
and
by
the
use
of
the
word
Evolution
twice
in
his
subsequently
published
(1866)
paper
that
summarized
both
talks.
Sadly,
the
converse
is
not
true,
despite
the
fact
that
Darwin's
fi
rst
cousin,
Francis
Galton,
wrote
to
him
in
1875
and
1876,
asking
him
to
grow
peas
to
test
Mendel's
laws.
Galton
also
recommended
that
Darwin
read
W.
0.
Focke's
Pflanzenmischlinge,
which
was
published
in
1881.
The
many
words
used
by
Darwin
in
the
fi
rst
edition
of
his
seminal
"On
the
Origin.
.
."
ignited
as
much
opposi-
tion
and
confusion
as
any
prior
eureka
in
biology.
Bate
-
son,
Mendel's
staunchest
and
most
verbal
protagonist
in
England,
used
Mendelism
to
bludgeon
natural
selection
acting
on
generationally
accrued,
infinitesimal
changes,
championing
instead
"sports",
mutations
of
major
devel-
opmental
effect
(e.g.,
"homoeotic"
ones,
Bateson
1894)
as
evolution's
raw
material.
It
was
not
until
1918
that
Fisher
reconciled
Mendelian
and
Galtonian
inheritance
thereby
initiating
the
genetic
study
of
evolution
making
genetics
and
descent
biologically
compatible
and
complementary
disciplines.
That
was
almost
60
years
after
"The
Ori-
gin.
.
."
Darwin
lacked
genetical
and
mathematical
foun-
dations
to
explain
descent
with
modification;
he
was
(in
our
opinion)
too
prolix,
occasionally
inconsistent,
con-
fused,
and
contradictory,
postulating
at
last
(again
after
Hippocrates)
pangenesis
and
ending
up
a
Lamarckian
2
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
J.
M.
Opitz
&
D.
W.
Bianchi
Invited
Commentary
(someone
whom
he
had
damned
roundly
on
previous
occasion).
By
contrast,
Mendel
was
concise,
distilling
the
essence
of
his
evidence
and
argument
into
just
47
pages
with
far
fewer
words
than
Darwin
required
to
make
his
point.
When
Mendel's
efforts
to
have
the
respected
botanist
Karl
Wil-
helm
von
NZgeli
at
the
University
of
Munich
repeat
his
Pi
-
sum
work
failed
(Correns
1905)
and
ended
in
utter
frustration,
he
went
on
with
life
and
work
as
the
abbot
of
his
community
tentatively
confident
that
"his
time
would
[still]
come."
When
it
came,
16
years
after
his
death
and
34
years
after
the
publication
of
Versuche,
it
occasioned
ini-
tial
amazement,
immediate
acceptance
and
the
kind
of
intellectual
joy
and
pleasure
attendant
on
and
reserved
for
only
those
few
truly
great
eurekas
of
Western
Biology
effec-
tive
in
reshaping
our
worldview,
akin
to
the
joy
occasioned
by
a
great,
beautiful,
unexpected
gift.
Not
just
something
given
and
taken
for
granted
as
self-evident,
but
a
stimulus
so
powerful
in
botany,
zoology,
and
medicine
as
to
occa-
sion
an
intellectual
revolution
akin
to
and
equal
in
effect
to
Darwinism
(Iltis
1924).
Iltis
put
it
nicely:
Mendelism
is
the
atomic
theory
of
life
(translation).
Thus,
when
Arnold
Lang
(1914)
of
Zurich
made
a
summary
of
all
of
Mendelism
in
zoology
up
to
the
beginning
of
the
First
World
War,
he
required
no
less
than
892
pages
for
Part
I;
we
do
not
know
if
Part
II
was
ever
published.
In
retrospect,
it
seems
hardly
believable
that
such
a
huge
amount
of
biology
involving
humans,
animals,
and
plants
wanted
to
and
needed
to
be
subsumed
under
the
head
of
Mendelism
so
rapidly
after
its
startling
birth
and
establishment
as
a
biological
discipline.
Mendel:
Morphologist
and
Mathematician
Mendel
not
only
observed
natural
phenomena,
as
a
well
-
trained
mathematician
and
physicist,
he
applied
mathe-
matical
theories
to
interpret
his
biological
experiments
and
to
plan
new
ones.
Iris
Sandler
(2000)
was
not
far
off
when
she
emphasized
the
concept
of
development
(Entwicklung)
in
Mendel's
paper,
except
that
she
still
confuses
(as
do
Stern
and
Sherwood
1966)
Entwicklung,
the
biological
entity,
with
Entwicklung
the
mathematical
entity.
As
the
late
C.W.
Cotterman,
then
also
at
the
University
of
Wisconsin,
made
clear
at
one
time
(Fig.
3):
".
.
.in
this
instance
it
seems
clear
that.
. .
[A]
poly-
nomial
(Polynom)
raised
to
a
power
(Potenz)
n
is
said
to
be
expanded
(entwickelt)
to
yield
a
series
(Reihe)
of
terms
(Glie-
der)
which
can
then
be
called
Entwickelungsreihe
[sic]
(i.e.,
(A
+
b)
2
=
A
2
+2Ab
+
b
2
).
Accordingly,
I
[Cotterman]
have
translated
this
as
a
'series
expansion.'"
1
And
in
working
on
a
renewed
translation
of
Mendel's
Versuche
from
the
ori-
ginal,
our
preference
will
be
"polynomial
expansion",
exactly.
Iris
Sandler's
confusion
is
no
different
from
that
of
William
Bateson
(1902)
and
the
Royal
Horticultural
Society
when
they
translated
this
as
"developmental
row,"
Ent-
wicklungsreihe
having
no
meaning
in
German
biology.
Near
the
beginning
of
the
19th
century
Goethe
(and
Burdach
in
1800)
introduced
the
term
Morphologic
into
biology
as
science
of
the
form,
formation,
transformation,
and
(after
Meckel
1812)
the
malformation
of
living
organisms
(Opitz
2004).
Form:
anatomy,
zootomy;
formation:
embryogenesis,
embryology;
transformation:
evolution
(after
the
Fr.
transformisme).
With
respect
to
malformation
(Missbildung):
Meckel
said
(Opitz
et
al.
2006)
they
were
"not
contrary
to
nature,"
and
like
their
then
normal
anatomical
counterparts,
were
quite
limited
in
type
and
range
since
"nature
is
not
infinitely
variable,"
being
merely
arrests
of
normal
development
or,
what
later
came
to
be
called
atavisms
(Darwin's
reversions),
abnor-
mal
in
humans,
normal
in
other
species.
This
limitation
is
now
referred
to
as
"developmental
constraint."
This
view
of
nature,
whether
philosophical
or
strictly
factual,
became
so
universally
understood
and
entrenched
in
European
thought
in
the
19th
century
as
to
make
its
explicit
invocation
as
methodological
approach
to
any
and
all
work
in
biology
redundant.
Mendel
worked
on
form
(tall,
short;
round,
wrinkled,
etc.)
and
its
formation
in
the
hybrids,
as
practical
an
approach
to
botanical
study
and
the
exploration
of
plant
development
as
that
of
his
father,
an
expert
fruit
tree
horticulturalist,
and
that
of
his
mother,
a
professional
gardener's
daughter
who
loved
ornamental
fl
owers.
Thus,
Mendel
did
not
have
to
mention
morphology
explicitly.
His
presentation
made
it
abundantly
clear
that
morphology
was
the
epistemologi-
cal
basis
of
his
botanical
and
later
apicultural
investiga-
tions.
In
part,
this
view
of
nature
(Merz,
1904-1912,
facsimile
1976)
was
acquired
"osmotically"
through
an
8
-
year
contact
with
his
in-house
Augustinian
brother
and
friend,
the
historian
and
philosopher
Fr.
Franz
Theodor
Bratranek,
a
confidant
of
Goethe's
daughter-in-law
and
her
sons
(Iltis
1924),
with
access
to
the
Weimar
Goethe
archives
and
an
editor
of
Goethe
papers.
Thus,
it
can
be
concluded
that
in
the
Altbrfinn
convent
at mid-century
Goethe
and
morphology
permeated
the
intellectual
atmo-
sphere
as
completely
as
oxygen
does
the
air.
And
Men-
del's
initial
approach
to
a
causal
analysis
of
development
in
plants
arose
out
of
very
real
morphology,
not
out
of
his
imagination
(Di
Trocchio
1991),
even
less
out
of
a
preconceived
notion
of
plant
genetics.
Whence
Mendel?
On
17
April
1850,
when
he
was
28
years
old,
Mendel
submitted
a
brief
autobiographical
note
together
with
other
documents
to
the
examination
commission
for
high
school
teachers
in
Vienna
(Iltis
1924).
It
breathes
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
3
Invited
Commentary
J.
M.
Opitz
&
D.
W.
Bianchi
The
word
Ehtwickelungsreihe
which
is
used
four
times
in
Mendel's
1866
paper,
as
well
as
The
word
Entwickelungsreihe
in
the
above
passage
is
one
which
Mendel
uses
4
times
in
his
famous
paper,
where
it
was
translated
by
Bateson
( )
and
the
Royal
Horticultural
Society
of
London
as
"developmental
series."
The
word
Entwicketung
waSralmost
,,
A
117
3
14.
'he
r
5
"44
11,4
"development"
if
the
application
is
a
biological
one.
But
in
this
instance
it
seems
cleli
o
that
44mirelplormanflo
7
rlimit
eertivemeakesi,ear.
A
polynomial
[Polynom]
raised
to
a
A
power
[Potenz]
n
is
said
to
be
expanded
[entwickelt]
to
yield
a
series
(htihel
of
ski.
terms
[Glieder],
which
can
A
he
called,Entwicketungsreihe.
Accordingly,
I
have
translated
this
as
a
"seriellexpansion."
The
word
Entwicketungsreihe
in
the
above
passage
is
one
which
Mendel
used
4
times
in
his
1866
paper,
where
it
was
translated
as
"developmental
series"
by
Bateson
and
the
Royal
Horticultural
Society
of
London
014,ke
Om)
d
o
a
ate,
gat
c!;tv
3(Id
4
T
arte
(Ae)
doer
cize:4
e.
.44<tiov
646(
j
,
er
4
';'
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i'eariz*
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1146
fri
Yi1R101.Af,11)
"140001
r
CWAViart
l
WEL
il
AIE.SRENial
DleOm
fi
ti/A-170
,16
EAIntl
Mendel's
humility
and
honesty.
Thereafter,
Mendel
did
not
indulge
in
autobiographical
documentation.
How
one
wishes
that
Mendel's
two
physician
nephews
had
undertaken
at
least
an
oral
history
while
their
uncle
was
living
and
they
were
in
frequent
contact
with
him.
Men-
del
made
their
education
possible
in
loving
gratitude
to
his
younger
sister
who
had
gladly
relinquished
her
dowry,
or
part
of
it,
so
that
Mendel
could
complete
his
own
(education).
In
1902,
when
Mendel
would
have
been
80,
a
memorial
tablet
was
dedicated
in
Heinzendorf
(HynCice,
Northern
Moravia),
Mendel's
birth
village,
and
affixed
to
the
local
fi
re
station,
where
he
had
orga-
nized
a
fi
re
brigade.
The
memorial
address
on
that
occasion
was
given
by
Dr.
Alois
Schindler,
the
older
of
Mendel's
two
nephews
whom
he
supported
during
medical
school.
This
necessarily
condensed
address
(Schindler
1902)
was
printed
privately
and
was
a
modest
beginning
of
the
Mendel
historiography
that
continues
to
this
day.
This
historical
process
was
severely
impaired
at
Figure
3.
Note
prepared
by
the
late
Charles
W.
Cotterman,
then
of
the
University
of
Wisconsin
on
the
term
Entwicklungsreihe
used
by
Mendel.
Date
?
Opitz
papers.
its
beginning
by
Mendel's
successor
as
abbot
who
destroyed
virtually
all
of
Mendel's
papers
after
his
death.
On
the
whole
though,
most
historians
of
Mendelism
initially
rely
on
Hugo
Iltis
of
Briinn,
whose
biography
of
Mendel
(1924)
was
thoroughly
researched
and
beautifully
written,
includ-
ing
an
exacting
analysis
of
Mendelism
to
that
date.
Dr.
Iltis
tells
charming
stories
of
Mendel
as
animal
lover,
for
all
except
snakes,
who
kept
a
fox
tied
up
in
the
morning
and
released
at
night,
and
who
made
a
startling
acquaintance
with
his
future
pet
hedgehog
who
had
bedded
down
for
the
night
in
one
of
his
boots.
The
two
monographs
of
Orel
(1984,
1996)
are
indis-
pensable
for
Mendel's
studies;
more
recently
Weiling
(1991),
published
a
summary
of
his
decades
-long
stud-
ies
of
personal
and
professional
aspects
of
Mendel's
life.
When
he
became
abbot,
Mendel
must
have
had
occa-
sional
moments
of
utter
astonishment
at
the
contrast
between
his
early
life
as
a
peasant's
son,
whose
father
still
labored
under
the
corvee
and
whose
incapacitation
while
4
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
J.
M.
Opitz
&
D.
W.
Bianchi
Invited
Commentary
felling
trees
in
the
lord's
forest
put
a
severe
crimp
on
the
continuation
of
Mendel's
education
and
his
present,
rela-
tively
opulent
life
as
prelate
of
a
wealthy
monastery.
His
attitude
toward
very
hard
work
on
his
own
and
the
lord's
land
never
left
Mendel,
who
did
not
allow
himself
a
moment
of
rest
after
beginning
monastic
life,
always
giving
the
very
best
of
himself
as
educator
(physics,
mathematics,
natural
history,
German,
Latin,
and
Greek),
as
practical
horticulturist,
investigator
of
plant
morphology,
meteorol-
ogist,
astronomer,
member
of
the
Moravian
legislature,
administrator
of
the
wealthiest
and
most
renowned
monas-
tery
of
Moravia
and
of
the
mortgage
bank
of
Moravia,
member
of
some
two
dozen
learned
societies
and
associa-
tions,
patron
of
the
arts,
and
benefactor
of
the
poor.
Mendel
Falsified?
It
would
be
naive
to
think
that
the
data
in
his
Versu-
che
were
the
only
ones
available
to
Mendel
for
his
pre-
sentation
or
manuscript
in
1865/6.
Mendel
was
an
extremely
hard
worker
and
it
can
be
assumed
without
risk
of
contradiction
that
he
performed
more,
perhaps
many
more
experiments
than
necessary
to
make
his
point.
As
an
outstanding
teacher,
renowned
for
his
didactic
skills,
he
evidently
used
great
care
in
sorting
his
material
to
make
as
convincing
a
case
as
possible.
Nowhere
in
the
records
from
over
a
half
century
have
we
ever
come
across
even
a
faint
hint
questioning
Mendel's
probity
or
honesty.
This
is
not
to
be
assumed
just
because
he
was
a
priest,
but
it
was
known
and
universally
acknowledged
by
his
contemporaries
that
Mendel
was
the
very
essence
of
integrity,
meriting,
after
all,
presidency
of
a
bank.
Also,
we
must
not
make
the
mistake
of
applying
pres-
ent
day
standards
of
data
analysis
and
presentation
to
the
style
and
standards
of
the
1860s;
Mendel
did
not
write
for
the
editors
of
Nature.
The
difference
is
of
the
essence
and
it
is
cultural,
not
a
failure
to
correctly
apply
Chi
-
squared
tests,
which
after
all
were
not
developed
until
after
Mendel's
death.
Mendel's
data
were
presented
not
to
deceive,
but
to
clarify
and
to
teach.
Furthermore,
his
writ-
ings,
especially
to
von
Na:geli
as
early
as
1866,
suggest
that
he
was
asking
for
others
to
validate
his
experimental
work.
Fisher
(1936)
concluded:
"There
can,
I
believe,
now
be
no
doubt
whatever
that
his
report
is
to
be
taken
entirely
lit-
erally,
and
that
his
experiments
were
carried
out
in
just
the
way
and
much
in
the
order
that
they
are
recounted."
But:
".
.
.the
data
of
most,
if
not
all,
of
the
experiments
have
been
falsified
so
as
to
agree
closely
with
Mendel's
expecta-
tions."
Sewall
Wright's
(1966)
reaction:
"The
most
serious
evidence
for
fraud
by
Mendel,
presented
by
Fisher,
is
the
very
close
agreement
to
a
ratio
of.
.
.."
Wright
redid
(some
of)
Fisher's
calculations,
agrees
with
his
results,
but
cautions:
"I
do
not
think
that
Fisher
allows
enough
for
the
cumulative
effect
on
X
2
of
a
slight
subconscious
tendency
to
favor
the
expected
result
in
making
tallies.
Mendel
was
the
fi
rst
to
count
segregants
at
all.
It
is
rather
too
much
to
expect
that
he
would
be
aware
of
the
precautions
now
known
to
be
necessary
for
com-
pletely
objective
data.
Anyone
who
doubts
the
difficulty
in
making
repeatable
counts
should
read
chapter
5
in
Pearl's
Introduction
to
Medical
Biometry.
He
reports
an
experiment
in
which
15
trained
observers
obtained
extraordinary
differences
in
sorting
and
counting
the
same
532
kernels
of
corn.
Checking
of
counts
that
one
does
not
like,
but
not
of
others,
can
lead
to
systematic
bias
toward
agreement.
I
doubt
whether
there
are
many
geneticists
even
now
whose
data,
if
extensive,
would
stand
up
wholly
satisfactorily
under
the
X
2
test.
"Men-
del's
perplexing
ratios
of
round
x
angular
and
yel-
low
x
green.
.
.
would
hardly
have
been
reported
by
one
bent
on
fraud".
Wright
concludes:
"Taking
every-
thing
into
account,
I
am
confident,
however,
that
there
was
no
deliberate
effort
at
falsification."
Similarly,
Sturtevant
(1985):
"Perhaps
the
best
answer
with
which
Fisher
would
have
agreed
is
that
after
all,
Mendel
was
right."
Weiling
(1991,
1993/1994)
has
examined
this
issue
on
several
occasions
concluding
that
".
.
.since
it
was
determined that
the
statistical
model
underlying
the
Chi
square
test
is
inappropriate
for
the
conditions
underlying
Mendel's
segregation
data,
. . .
Fisher's
conclusion
does
not
apply."
Edwards
(1993):
"As
to
the
controversy
over
his
goodness
-of
-fit
fi
ndings
. .
.the
less
said
the
better.
The
only
people
on
whom
it
reflects
badly
are
those
writers
who
have
viewed
it
as
casting
doubt
on
the
scientific
integrity
of
either
Mendel
or
Fisher."
Conclusions
Assuming
the
sexuality
of
plants,
the
equivalence
of
pol-
len
and
ovule
in
the
production
of
offspring
and
the
self-
evident
transmission
of
plant
traits
over
generations
("inheritance"),
the
following
conclusions
may
be
drawn
from
elementary
Mendelism
involving
pairs
of
contrasting
traits
(Fig.
4).
Exponent:
Number
of
traits;
a
having
the
property
of
becoming
latent
("recessive")
in
the
hybrid
dominated
by
the
"domineering"
A
thus
designated
subsequently
the
dominant
(partner)
trait.
The
following
10
inferences
may
be
drawn
from
these
results:
1
Determinants
of
A
and
a
are
concrete
material,
partic-
ulate
entities.
. .
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
5
Invited
Commentary
J.
M.
Opitz
&
D.
W.
Bianchi
Hybrid
Crossbred
Progeny
A
and
a
Aa
Aa
x
Aa
AA
+
Aa
+
aA
+
aa
Phenotypic
ratio
3A
:
la
rlY
Aa
selfed
AA
Aa
aA
aa
Ratio
1
:
2
:
1
breed
true
hybrids
breed
true
Expansion
of
4,
1
1A
2
:
2Aa
la
g
(A
+
a)
2
binomial
Figure
4.
Succinct
summary
of
Mendelism
to
Mendel's
death
in
1884.
2
..
.
.which
are
apparently
assembled
in
(a
pairwise
man-
ner)
in
hybrids
without
merger
or
loss
of
material
integrity.
. .
3
..
.
.as
shown
by
their
intact
individual
reappearance
("segregation",
Bateson
and
Saunders
1902)
in
the
germ
cells
and
4
..
.
.the
reappearance
of
the
recessive
trait
in
the
prog-
eny
of
the
hybrids.
5
The
determinants
of
A
and
a
must
be
equivalent
in
structure
and
function
to
judge
from
their
1:
1
occur-
rence
in
the
offspring
of
the
hybrids.
. .
6
..
.
.regardless
whether
carried
by
pollen
or
ovules.
7
Determinants
of
A
and
a
are
the
morphological
units
of
function
affecting
the
development
of
the
character
traits;
as
such.
. .
8
..
.
.they
may
be
pleiotropic
(Plate
1910)
and
. . .
9
..
.
.have
to
be
taken
into
account
of
any
trait
affecting
evolution
since.
. .
10
..
.
.they
may
be
responsible,
wholly
or
in
part,
for
those
traits
deemed
"desirable"
by
the
breeder,
thus
leading
in
the
short
term
to
the
appearance
of
"new"
races,
breeds
and
cultivars
by
artificial
selection.
Fisher
(1936),
referring
to
Focke's
Pflanzenmischlinge
which,
like
Nageli,
completely
missed
the
epochal
implica-
tions
of
Mendel's
work,
stated
".
.
.the
learned
author
(Focke)
having
overlooked,
in
his
chosen
fi
eld,
experimental
researches
conclusive
in
their
results,
faultlessly
lucid
in
pre-
sentation,
and
vital
to
the
understanding
not
of
one
problem
of
current
interest,
but
of
many"
(italics
added)".
In
retrospect,
no
one
can
disagree
with
this
assessment
of
Mendel's
work.
From
the
surviving
records,
the
testimony
of
contempo-
raries
and
the
evidence
collected
by
historians
closest
to
Mendel
in
time
(but
all
after
1900)
it
seems
safe
to
con-
clude
that
above
all
Mendel
was
a
"good"
man
in
the
best
meaning
of
the
word.
Not
just
by
ecclesiastical
standards,
Mendel
being
only
one
of
many
abbots
in
Western
Catholi-
cism,
but
fi
rst
of
all
by
the
astonishing
outpouring
of
grief
at
his
death,
for
example,
by
his
former
students,
fellow
pedagogues
and
by
the
poor
citizens
of
the
town.
And
what
a
legacy
for
an
abbot
(!)
to
bequeath
to
humanity,
no
hymns,
no
theological
treatises,
no
ser-
mons,
no
record
of
particular
sanctity
(indeed
the
opposite
in
his
obstinate
refusal
to
pay
the
church
tax).
Rather,
a
brief
47
-page
monograph
on
the
progeny
of
plant
hybrids,
work
conceived
and
executed
over
10
years
apparently
without
preconceived
notions,
and
confined
strictly
to
what
we
now
call
phenotypes
and
the
behavior
of
their
formbildenden
Elemente
in
the
germ
cells.
This
astonishing
epistemological
restraint
made
it
so
easy
for
his
successors,
for
example,
De
Vries,
Correns,
Tschermak,
Bateson.
.
.to
take
the
next
step
to
"genotype".
We
agree
with
Olby
(1966,
1979,
1991)
that
Mendel
did
not
set
out
to
discover
Mendelism,
or
to
plan
his
laborious
experiments
accordingly,
or
to
"falsify"
his
results
so
as
to
best
fi
t
expectations
(Fisher
1936).
We
differ
only
slightly
from
Fisher
in
that
Mendel
probably
did
not
stop
counting
when
he
had
made
his
point,
more
or
less
exactly,
but
rather
selected
the
best
results
from
many.
During
the
dreadful
time
of
war
after
his
presentations,
the
occupation
of
Briinn
by
some
50,000
Prussians
in
early
1866,
and
the
ensuing
cholera
epidemic
and
food
shortages
Mendel
had
some
time
to
reflect
on
his
manuscript
before
its
publica-
tion
in
1866.
He
summarized
his
results
in
the
didactically
clearest
manner,
not
to
deceive
but
to
impress
as
parsimo-
niously
as
possible.
Additionally,
his
use
of
comparative
mathematics
to
interpret
biology
was
ahead
of
its
time,
hav-
ing
important
parallels
today
in
the
use
of
computational
biology
to
analyze
big
datasets.
Mendel
was
a
geneticist
and
genomicist
malgre
lui.
In
the
150
years
since
Mendel's
oral
presentations,
we
have
taken
the
inferences
drawn
from
his
work
so
far
as
to
begin
an
inventory
of
the
genes
in
LUCA,
an
organism
that
lived
some
3.6
billion
years
ago.
LUCA
became
the
ancestor
of
all
living
(and
extinct)
pro-
and
eukaryotes
(Goldman
et
al.
2013;
on
LUCApedia).
These
astonishing
advances
and
capabilities
notwithstanding,
Mendel
would
probably
be
content
to
have
made
a
begin
-
6
©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
J.
M.
Opitz
&
D.
W.
Bianchi
Invited
Commentary
ning
at
the
phenome,
or
rather
at
phenomics,
genomics
having
become
one
of
those
unanticipated
consequences
arising
from
the
work
of
an
astute
man
far
ahead
of
his
times.
Conflict
of
Interest
None
declared.
Note
1
Cotterman,
a
colleague
and
friend
of
John
M.
Opitz
in
Madison
for
18
years
before
his
death,
was
not
only
an
expert
germanist
but
also
mathematician,
the
equal
of
Mendel
and
for
a
while,
the
world
pioneer
and
greatest
authority
on
combinatorics
and
path
coefficients
applied
to
humans
-
specifically
immunogenetics.
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©
2015
The
Authors.
Molecular
Genetics
&
Genomic
Medicine
published
by
Wiley
Periodicals,
Inc.
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