-
1
MC
B14
0 01
-29-
07 1
Gen
etic
s –
the
narr
ativ
e so
far
1.M
ende
l (18
53-6
6):
1.H
ered
ity o
pera
tes
via
parti
cles
of i
nfor
mat
ion
(=ge
nes)
.2.
For a
ny tr
ait,
two
parti
cles
(=al
lele
s of
a g
ene)
com
e to
geth
er in
an
orga
nism
, but
then
sep
arat
e in
to g
amet
es, o
ne p
er g
amet
e.3.
This
pro
cess
occ
urs
inde
pend
ently
for p
artic
les
(gen
es) f
or d
iffer
ent
traits
.2.
Häc
kel,
Wei
ssm
ann,
Fle
mm
ing,
Bov
eri (
1850
-190
0):
1.Th
e nu
cleu
s of
the
cell
is th
e or
gane
lle th
at is
repo
nsib
le fo
r her
edity
(H
ääck
el, B
over
i), a
nd s
peci
fic k
inds
of c
ells
(the
gam
etes
) tha
t are
se
para
te a
nd d
istin
ct fr
om th
e re
st o
f the
bod
y, a
re re
spon
sibl
efo
r ge
nera
ting
prog
eny
(Wei
ssm
an).
2.In
side
the
cell,
lie
the
chro
mos
omes
(Fle
mm
ing)
that
sep
arat
e in
to
daug
hter
cel
ls v
ia m
itosi
s.3.
Pro
per d
evel
opm
ent o
f an
orga
nism
requ
ires
a sp
ecifi
c se
t of a
pa
rticu
lar k
ind
of c
hrom
osom
e (B
over
i).3.
Sutto
n (a
nd C
anno
n) –
1902
.1.
Ther
e is
a s
triki
ng c
onco
rdan
ce b
etw
een
the
beha
vior
of M
ende
l’spa
rticl
es a
nd c
hrom
osom
es d
urin
g m
eios
is.
MC
B14
0 01
-29-
07 2
Gen
etic
s –
the
narr
ativ
e to
com
e1.
Mor
gan
and
Brid
ges
–ge
nes
lie o
n ch
rom
osom
es (1
910-
1915
).2.
Bat
eson
and
Stu
rteva
nt (1
906-
15) –
som
e ge
nes
appe
ar to
be
phys
ical
ly
linke
d; th
e na
ture
of t
his
linka
ge is
the
follo
win
g: g
enes
are
arr
ange
d on
a
chro
mos
ome
in a
line
ar o
rder
, at p
artic
ular
dis
tanc
es fr
om e
ach
othe
r.3.
McC
linto
ck a
nd S
tern
(193
0) –
gene
tic re
com
bina
tion
occu
rs w
hen
hom
olog
ous
chro
mos
omes
can
exc
hang
e pa
rts.
4.B
eadl
e an
d Ta
tum
(194
6) –
som
e ge
nes
affe
ct th
e de
velo
pmen
t of t
raits
by
enc
odin
g bi
oche
mic
al fu
nctio
ns (“
one
gene
= o
ne e
nzym
e”).
5.M
ulle
r (19
27) –
the
gene
can
be
purp
osef
ully
mut
ated
.6.
Ben
zer –
the
gene
itse
lf ca
n be
spl
it in
to s
mal
ler u
nits
.7.
Stu
dyin
g th
e m
olec
ular
mak
eup
of li
fe v
ia:
1.Th
e ge
netic
scr
een
–us
e of
pur
pose
ful m
utag
enes
is to
iden
tify
gene
s re
quire
d fo
r the
dev
elop
men
t of v
ario
us tr
aits
.2.
Map
ping
by
linka
ge a
nd a
ssoc
iatio
n m
appi
ng –
the
stud
y of
“sim
ple”
and
“c
ompl
ex” t
raits
by
anal
ysis
of p
edig
rees
and
pop
ulat
ions
.
MC
B14
0 01
-29-
07 3
Wha
t is
bein
g om
itted
fo
r lac
k of
tim
e1.
The
redi
scov
ery
of M
ende
l’s la
ws
by
Cor
rens
, Tsc
herm
ak, a
nd d
e V
ries.
2.Th
e fin
ding
–by
Cue
not a
nd b
y C
astle
–th
at M
ende
l’s la
ws
also
app
ly to
m
amm
als,
suc
h as
mic
e an
d gu
inea
pi
gs.
MC
B14
0 01
-29-
07 4
Willi
am B
ates
on
“…he
priv
atel
y su
bsid
ized
his
sm
all b
ook,
Men
del’s
Prin
cipl
es
of H
ered
ity: A
Def
ence
and
he
sent
cop
ies
to a
ll of
the
lead
ing
stud
ents
of h
ered
ity to
mak
e su
re
that
Men
del w
ould
not
suf
fer
anot
her 3
5 ye
ars
of n
egle
ct.”
Car
lson
Men
del’s
Leg
acy
-
2
MC
B14
0 01
-29-
07 5
Mod
ified
Men
delia
n ra
tios
epis
tasi
s
MC
B14
0 01
-29-
07 6
“Rep
ulsi
on a
nd c
oupl
ing”
linka
ge
MC
B14
0 01
-29-
07 7
Bat
eson
190
2M
CB
140
01-2
9-07
8B
ates
on 1
902
αλληλος
= "e
ach
othe
r"
-
3
MC
B14
0 01
-29-
07 9
Fig.
3.7
MC
B14
0 01
-29-
07 1
0
Thom
as H
unt M
orga
n, th
e fir
st n
ativ
e-bo
rn A
mer
ican
to w
in th
e N
obel
Priz
e,
foun
der o
f mod
ern
gene
tics
Brid
ges
Mul
ler
Mor
gan
Stu
rteva
nt
MC
B14
0 01
-29-
07 1
2
Mor
gan
et a
l. 19
15
“As
will
be s
how
n no
w, c
erta
in fa
ctor
s fo
llow
th
e di
strib
utio
n of
the
X c
hrom
osom
e an
d ar
e th
eref
ore
supp
osed
to b
e co
ntai
ned
in th
em.”
Em
phas
is m
ine
–fd
u.
↓G
enes
lie
on c
hrom
osom
es
-
4
MC
B14
0 01
-29-
07 1
3
“The
sup
posi
tion
that
par
ticle
s of
ch
rom
atin
, ind
istin
guis
habl
e fro
m
each
oth
er a
nd in
deed
alm
ost
hom
ogen
eous
und
er a
ny
know
n te
st, c
an b
y th
eir m
ater
ial
natu
re c
onfe
r all
the
prop
ertie
s of
lif
e su
rpas
ses
the
rang
e of
eve
n th
e m
ost c
onvi
nced
mat
eria
lism
.”
Bat
eson
, W. (
1916
) The
m
echa
nism
of M
ende
lian
here
dity
(a
revi
ew).
Sci
ence
, 44,
536
-543
.
MC
B14
0 01
-29-
07 1
4
MC
B14
0 01
-29-
07 1
5
A p
robl
em a
nd a
sol
utio
n“T
he v
alue
and
util
ity o
f any
exp
erim
ent…
” (M
ende
l)“W
hat w
as n
eede
d to
ope
n up
gen
etic
s to
new
ph
enom
ena
was
an
orga
nism
that
bre
d ra
pidl
y,
prod
uced
lots
of p
roge
ny, a
nd w
as in
expe
nsiv
e to
mai
ntai
n” (C
arls
on)
“Fru
it fli
es c
an b
e ra
ised
on
a m
ixtu
re o
f cor
n m
eal,
yeas
t, su
gar,
and
agar
. Flie
s co
mpl
ete
thei
r life
cy
cle
from
ferti
lizat
ion
to e
mer
genc
e of
the
adul
t fly
in 1
0 da
ys. A
fem
ale
can
prod
uce
3,00
0 pr
ogen
y in
her
life
time.
A s
ingl
e m
ale
can
sire
w
ell o
ver 1
0,00
0 of
fspr
ing.
” (H
artw
ell)
MC
B14
0 01
-29-
07 1
6
(aka
chr
. 1)
Not
e: n
o cr
ossi
ng o
ver i
n m
ale
mei
osis
!
-
5
MC
B14
0 01
-29-
07 1
8
Mor
gan
and
Dro
soph
ila(g
o B
ears
)M
orga
n w
as n
ota
gene
ticis
t by
train
ing
(he
was
an
embr
yolo
gist
), an
d he
w
as n
otth
e fir
st o
ne to
use
Dro
soph
ila fo
r pur
pose
s of
gen
etic
rese
arch
(C
astle
was
).“O
ne o
f the
baf
fling
pro
blem
s of
bre
eder
s in
pre
-Men
delia
n da
ys h
ad
been
the
effe
cts
of in
bree
ding
and
cro
ssbr
eedi
ng. W
hat t
hese
wer
e w
as
a m
uch-
deba
ted
ques
tion.
We
set o
ut to
giv
e it
an e
xper
imen
tal t
est a
ndfo
und
read
y to
han
d a
rapi
dly
bree
ding
littl
e fly
, Dro
soph
ila, b
eing
cu
lture
d in
the
labo
rato
ry b
y a
grad
uate
stu
dent
as
embr
yolo
gica
l m
ater
ial.
This
, he
told
us,
wou
ld c
ompl
ete
a ge
nera
tion
with
in a
fortn
ight
. (C
harle
s W
oodw
orth
, pro
f ent
omol
ogy
at U
C B
erke
ley)
. … W
e b
egan
cultu
ring
the
fly o
n pu
lped
Con
cord
gra
pes,
but
this
gav
e us
poo
r res
ults
as
man
y of
the
larv
ae w
ould
get
dro
wne
d an
d th
en o
ur p
opul
atio
n st
atis
tics
wer
e no
goo
d. A
s gr
apes
bec
ame
out o
f sea
son,
we
tried
oth
er
fruits
, and
fina
lly h
it th
e ja
ckpo
t in
bana
nas.
…Th
e co
nclu
sion
dra
wn
[from
our
stu
dies
] was
that
inbr
eedi
ng re
duce
s ve
ry s
light
ly th
epr
oduc
tiven
ess
of D
roso
phila
. … T
his
was
not
a c
oncl
usio
n of
wor
ld-
shak
ing
impo
rtanc
e. T
he im
porta
nt o
utco
me
of th
is in
vest
igat
ion
was
that
it
calle
d to
Mor
gan’
s at
tent
ion
a ne
w s
ourc
e of
mat
eria
l for
exp
erim
enta
l st
udy
not s
ubje
ct to
the
limita
tions
of s
low
-bre
edin
g la
bora
tory
mam
mal
s.”
WE
Cas
tle (p
rof g
enet
ics
UC
Ber
kele
y) T
he B
egin
ning
s of
Men
delis
m in
A
mer
ica
–in
Gen
etic
s in
the
20th
Cen
tury
, p. 7
3.
MC
B14
0 01
-29-
07 1
9
Toug
h ea
rly g
oing
“…M
orga
n ha
d be
en w
orki
ng
on fr
uit f
lies
for a
t lea
st tw
o ye
ars
befo
re h
e fo
und
his
mos
t sig
nific
ant m
utat
ion,
a
whi
te-e
yed
fly. F
or th
is n
ew
appr
oach
, Mor
gan
was
his
ow
n fir
st s
tude
nt. H
e br
ed
the
flile
s fo
r tw
o ye
ars
with
out a
ssis
tanc
e. …
He
poin
ted
to th
e sh
elve
s w
ith
flies
and
[sai
d] th
at h
e ha
d w
aste
d tw
o ye
ars
and
had
gotte
n no
thin
g fo
r his
wor
k.”
MC
B14
0 01
-29-
07 2
0
“May
191
0 w
as w
hen
the
revo
lutio
n be
gan.
Mor
gan
foun
d a
whi
te-e
yed
mal
e ru
nnin
g ar
ound
in o
ne b
ottle
.”
-
6
MC
B14
0 01
-29-
07 2
1
Gre
at o
peni
ng p
assa
ges
in th
e hi
stor
y of
the
Eng
lish
lang
uage
“It is
a tr
uth
univ
ersa
lly a
ckno
wle
dged
that
a
sing
le m
an in
pos
sess
ion
of a
goo
d fo
rtune
m
ust b
e in
wan
t of a
wife
.”
“Whe
n C
arol
ine
Mee
ber b
oard
ed th
e af
tern
oon
train
for C
hica
go, h
er to
tal o
utfit
co
nsis
ted
of a
sm
all t
runk
, a c
heap
imita
tion
allig
ator
-ski
n sa
tche
l, a
smal
l lun
ch in
a
pape
r box
, and
a y
ello
w le
athe
r-sn
ap p
urse
, co
ntai
ning
her
tick
et, a
scr
ap o
f pap
er w
ith
her s
iste
r’s a
ddre
ss in
Van
Bur
en S
treet
, an
d 4
dolla
rs in
mon
ey. I
t was
in A
ugus
t, 19
98. S
he w
as e
ight
een
year
s of
age
, br
ight
, tim
id, a
nd fu
ll of
the
illus
ions
of
igno
ranc
e an
d yo
uth.
”
MC
B14
0 01
-29-
07 2
2
MC
B14
0 01
-29-
07 2
3
Req
uire
d re
adin
g fo
r Mor
gan’
s S
cien
ce p
aper
1.Th
e pa
per i
tsel
f.2.
The
com
men
tary
to it
, writ
ten
by R
ober
t R
obbi
ns, t
hat p
rece
des
the
pape
r its
elf i
n th
e E
SP
-gen
erat
ed P
DF.
3.M
orga
n’s
own
desc
riptio
n in
nar
rativ
e fo
rm, p
p. 1
5-20
of h
is 1
915
book
(als
o an
E
SP
-pro
vide
d P
DF)
.
MC
B14
0 01
-29-
07 2
4
“The
Mec
hani
sm o
f M
ende
lian
Her
edity
” (19
15)
Thom
as H
unt M
orga
nA
lfred
Stu
rteva
ntH
erm
ann
Mul
ler
Cal
vin
Brid
ges
-
7
MC
B14
0 01
-29-
07 2
5M
CB
140
01-2
9-07
26
MC
B14
0 01
-29-
07 2
7
Not
hing
spe
cial
her
e.
Just
like
see
d co
lor i
n pe
as.
Nor
mal
Men
delia
n ra
tio (3
:1) –
but w
here
ar
e th
e w
hite
-eye
d fe
mal
es?!
!
MC
B14
0 01
-29-
07 2
8
-
8
MC
B14
0 01
-29-
07 2
9M
CB
140
01-2
9-07
30
Net
tie S
teve
ns, d
isco
vere
r of t
he
sex
chro
mos
omes
Net
tie S
teve
ns w
as o
ne o
f the
firs
t fem
ale
scie
ntis
ts to
mak
e a
nam
e fo
r her
self
in
the
biol
ogic
al s
cien
ces.
She
was
bor
n in
Cav
endi
sh, V
erm
ont.
Her
fam
ily
settl
ed in
Wes
tford
, Ver
mon
t. S
teve
ns' f
athe
r was
a c
arpe
nter
and
han
dym
an.
He
did
wel
l eno
ugh
to o
wn
quite
a b
it of
Wes
tford
pro
perty
, and
cou
ld a
fford
to
sen
d hi
s ch
ildre
n to
sch
ool.
Ste
vens
was
a b
rillia
nt s
tude
nt, c
onsi
sten
tly s
corin
g th
e hi
ghes
t in
her c
lass
es. I
n 18
96, S
teve
ns w
ent t
o C
alifo
rnia
to a
ttend
Lel
and
Sta
nfor
d U
nive
rsity
. She
gr
adua
ted
with
a m
aste
rs in
bio
logy
. Her
thes
is in
volv
ed a
lot o
f mic
rosc
opic
w
ork
and
prec
ise,
car
eful
det
ailin
g of
new
spe
cies
of m
arin
e lif
e. T
his
train
ing
was
a fa
ctor
in h
er s
ucce
ss w
ith la
ter i
nves
tigat
ions
of c
hrom
osom
al
beha
vior
.A
fter S
tanf
ord,
Ste
vens
wen
t to
Bry
n M
awr C
olle
ge fo
r mor
e gr
adua
te w
ork.
Th
omas
Hun
t Mor
gan
was
stil
l tea
chin
g at
Bry
n M
awr,
and
was
one
of h
er
prof
esso
rs. S
teve
ns a
gain
did
so
wel
l tha
t she
was
aw
arde
d a
fello
wsh
ip to
st
udy
abro
ad. S
he tr
avel
ed to
Eur
ope
and
spen
t tim
e in
The
odor
Bov
eri's
lab
at th
e Zo
olog
ical
Inst
itute
at W
urzb
urg,
Ger
man
y. B
over
i was
wor
king
on
the
prob
lem
of t
he ro
le o
f chr
omos
omes
in h
ered
ity. S
teve
ns li
kely
dev
elop
ed a
n in
tere
st in
the
subj
ect f
rom
her
sta
y.In
190
3, S
teve
ns g
ot h
er P
h.D
. fro
m B
ryn
Maw
r, an
d st
arte
d lo
okin
g fo
r a
rese
arch
pos
ition
. She
was
eve
ntua
lly g
iven
an
assi
stan
tshi
p by
the
Car
negi
e In
stitu
te a
fter g
low
ing
reco
mm
enda
tions
from
Tho
mas
Hun
t Mor
gan,
Edm
und
Wils
on a
nd M
. Car
ey T
hom
as, t
he p
resi
dent
of B
ryn
Maw
r. H
er w
ork
on s
ex
dete
rmin
atio
n w
as p
ublis
hed
as a
Car
negi
e In
stitu
te re
port
in 1
905.
MC
B14
0 01
-29-
07 3
1
The
usef
ul m
ealw
orm
“It is
per
fect
ly c
lear
that
an
egg
ferti
lized
by
a sp
erm
atoz
oon
cont
aini
ng th
e sm
alle
r he
rero
chro
mos
ome
prod
uces
a
mal
e, w
hile
one
ferti
lized
by
a sp
erm
atoz
oon
cont
aini
ng th
e la
rger
het
eroc
hrom
osom
e de
velo
ps in
to a
fem
ale.
”X
O is
mal
e an
d X
X is
fem
ale
(unt
il 19
16)
MC
B14
0 01
-29-
07 3
2
Edm
und
Wils
on –
argu
ably
, the
firs
t X
Y p
erso
n on
Ear
th“In
man
y sp
ecie
s of
inse
cts
ther
e ar
e tw
o cl
asse
s of
spe
rmat
ozoa
, equ
al in
nu
mbe
r, w
hich
in th
e ea
rly s
tage
s of
thei
r de
velo
pmen
t, di
ffere
vis
ibly
in re
spec
t to
the
nucl
ear c
onst
itutio
n; w
hile
ther
e is
bu
t one
cla
ss o
f egg
. … T
hat i
s to
say
, if
the
two
kind
s of
spe
rmat
ozoa
be
desi
gnat
ed a
s th
e X-
clas
s an
d th
e Y
-cl
ass,
resp
ectiv
ely;
the
eggs
are
all
of
the
X-cl
ass.
The
mal
e m
ay a
ccor
ding
ly
be d
esig
nate
d as
the
hete
roga
met
ic s
ex,
the
fem
ale
as th
e ho
mog
amet
ic.”
-
9
MC
B14
0 01
-29-
07 3
3
Dr.
Thom
as R
ied,
NC
I/NIH
:
SK
Y o
f hor
mal
hum
an c
ell
MC
B14
0 01
-29-
07 3
4
MC
B14
0 01
-29-
07 3
5M
CB
140
01-2
9-07
36
-
10
MC
B14
0 01
-29-
07 3
7M
CB
140
01-2
9-07
38
MC
B14
0 01
-29-
07 3
9M
CB
140
01-2
9-07
40
-
11
MC
B14
0 01
-29-
07 4
1
