Professional Documents
Culture Documents
C.
H.
WADDINGTON
How Animals
Develop
'l.L
lllustraied
L
tt^jf-f
CD
HOW
ANIMALS DEVELOP
C. H. Waddington,
M. A.
HARPER TORCHBOOKS
The Science Library
& Brothers
New York
Harper
HOW ANIMALS
New
DEVELOP
material Copyright
@ 1962
by C. H. Waddington
FOREWORD
In this book,
it
was impossible
to shirk
which
HOW
ANIMALS DEVELOP
C. H.
W.
CAMBRIDGE
1935
ACKNOWLEDGMENTS
For
made
is
to the
CONTENTS
PAGE
CIIAPflR
FOREWORD
I.
INTRODUCTION
I3
of
all
organization
young embryos
The
The
three
fundamental layers
II.
THE
BEGINNING
DEVELOPMENT
OF
birth
III.
26
heredity, virgin
4O
IV.
THE
"organization CENTRE*'
The technique
of
micro-surgery
is
61
The focus
Birds
integrated
THE
ADDITION
OF
DETAILS
78
VI.
VII.
93
Regeneration
102
How
embryos are
of function
in
development
INDEX
135
LIST OF ILLUSTRATIONS
PAGE
FIG.
1.
2.
3.
4.
5.
6.
7.
8.
9a.
26
2Q
29
3I
33
36
38
42
facing page 42
gb.
10.
11.
12.
13.
14.
15.
ON
THE
45
46
5I
52
54
57
HOW
ANIMALS DEVELOP
PAGE
FIG.
exchange
i6.
spemann's
17.
the newt
an organizer graft
AN organizer graft
experlment
IN
18.
in
IN
the newt
THE CHICK
65
68
facing page 70
19.
20.
21.
22.
23.
24.
25.
L.N.E.R.
26.
27.
28.
29.
facing page 93
IO7
IIO
112
facing page
1 1
INTRODUCTION
The Development of Animal Organization
we
organs,
find that
and
it,
an animal
in all of
them
and
is
so on. If
we
look
made up of different
there
is
something going
On
on
all
cell is
the time.
always
full
many
and pours out again into its surroundings the byproducts for which it has no use.
In a living organism these changes are not isolated
but are adjusted to one another so that the right
operations are carried out to produce the right
quantities of the various products.
It
is
because
14
HOW
we
ANIMALS DEVELOP
way
here
is
we must
look for
in
is
a "secret of life,"
among
it,
the causes
When
to
be organized
of processes
is
is
it
set
among
jobs
which
specializes
function.
the
in carrying
carburettor
^to
vaporize the
fuel,
another
still
and
so on.
apparatus
in all
^to
We
size.
to transport
it
it.
We
lungs in animals
INTRODUCTION
many
velopment of a
set
of specialized structures
is
The
de-
the
first
up a living organism.
To say that an animal is an organism means in
fact two things firstly, that it is a system made up
of separate parts, and secondly, that in order to
describe fully how any one part works one has to
Thus
it is
organism
is
The
fits
on
it is
to a pelvis
part of a leg,
if
an anatomist
finds
HOW
ANIMALS DEVELOP
about, but
we cannot
first
place.
as
amount of food-matter or
yolk,
bag of special
kernel or nucleus. As we shall
encloses a
little
protoplasm
is
the jelly-like
see,
it
looks.
But
it
at
animal
arises.
In the early
it
is
fluid
its
it,
or embryology, because
of finding out
how
the most
INTRODUCTION
1 *]
embryology remained
a purely descriptive science. The changes which
embryos go through as they develop are so many
and complicated that it took an enormous amount
of careful and painstaking work simply to describe
them. Scientists have always asked why the changes
occur; but only in the last fifty years or so have
they been able to perform experiments to try to find
centuries,
at
of an embryo
they work in
this stage a lung can develop quite independently of the heart,
but it cannot function to aerate the blood without the help of
^
Though
a heart.
HOW
ANIMALS DEVELOP
way
way any
part develops
is
is
available
is
is
The
con-
life,
It
INTRODUCTION
us.
The Similarity of
The
all
young Embryos
fillip
by
publication,
a bird or a
classified as
mammal,
while
still
later
it
animal, as
it
one of
its
series
is
more or
is
The
which
less in
it
belongs.
first
generalized ancestral
HOW
20
ANIMALS DEVELOP
vertebrate
stock.
theory.
Many
details
of
this
modification,
INTRODUCTION
larvae.
law
Haeckel's
When
of anything.
its
remote
rotted to
is
fish-like
mud on
not
strictly
an
human embryo
ancestors
is
explanation
developing,
and
why
We
evolutionary
change from
obviously easier, or so a
fish
into
man would
men,
it
is
think, to stick
HOW
22
ANIMALS DEVELOP
call this
to
it is
men
inventing,
do what
their
easier.
to
ancestral
embryos only
for
this sort
characters
short
of explanation
which appear
time,
eventually
in
dis-
appearing entirely.
The Three Fundamental Layers
INTRODUCTION
of animals had been investigated, and
23
it
had been
The
original
adult, but
no animals of
this
kind
The
cell-thick.
but
differs
which
lines
it, is
as a
little
HOW
24
surface
ANIMALS DEVELOP
it
first visible
indication
is
them
endoderrriy
and
mesoderm.
are the
is
We
The mesoderm
INTRODUCTION
century, and
on which
it
all
25
On
the whole,
it
was
difficult
cases,
and
in
is
usually the
first
is
blocked out.
when
the
main
CHAPTER
II
The
when
series
of processes by which
NUCLEUS
...
O*
T.
HEAD
CENTREPIECE
TAIL
Diagram
Fig.
I.
sperm
is
including the
tail
known
collectively as germ-cells,
nucleus.
(Fig.
unlike normal
i)
But in the
details
of their
is
cells, as
it
has to
27
is
grains
as
through the
of yolk,
cytoplasm, that
is
usually
lies
in
the
The
clear
human
is
about o
mm.
or a
The
sperm-cell
is still
:
can travel
process
essentially the
is
in the sperm.
same both
in the eggs
and
HOW
28
an ordinary
cell consists
membrane
filled
the
is
cell
membrane
if
made
of the nuclear
When
up a number of
the cell
many
of a bag
which
ANIMALS DEVELOP
is
killed
liquid contents
little solid
lumps,
dyes,
The
difference
is
this: that
Fig. 3.
Fig. 2.
29
The chromoDiagram
somes appear, already double, in the nucleus, (b) The cell
divides and one-half of each double chromosome goes to
each daughter cell
Fig. 2,
HOW
30
ANIMALS DEVELOP
istics
of the germ-cells.
The
ordinary body-cells,
which
all
31
it
number to half.
The daughter-cells of
their
MATURATION PERIOD
O
REDUCTION
DIVISION
REDUCTION
DIVISION
Qo
06
6606
PERIOD OP
DIFFERENTIATION
Diagram
it
is
ready,
and
HOW
32
ANIMALS DEVELOP
Fertilization^ Heredity^
At the end of
ready to
all this
carry
Virgin Birth
out
the
complicated
of
process
Fertilization
really
consists
of two
by the sperm
and sperm nuclei. It is
o^gg
it
of chromosomes
is
development of
An example
will
Men
are some-
some abnormality
33
it
is
the
therefore said to be
PARENTS
Ss
St
GERM-CELLS
CHILDREN
ISS
2 St
Iss
gene, which
is
recessive to
it.
When
the germ-cells
a quarter of
more
short-fingered adults,
and
HOW
34
ANIMALS DEVELOP
Hereditary factors of
Mendel
also
this
and he
inherited.
that time,
The
of the tgg.
happens.
We know
What
it
does
very
is
little
is
the activation
about
how
this
Now
we then
it
is
35
The most
called in science.
various
effective. It is
some-
eggs
may be
it
into very
weak
acid
some marine
is
altered.
In
if
all
the
cases
we
have very little idea why they give any results at all.
But the eggs treated in this way, since they have a
haploid set of chromosomes, can go on developing
quite normally. The adult which arises is smaller
than normal, and its cells are smaller than normal,
since they adjust their volume to that of the half-sized
nuclei. Some eggs normally develop without being
fertilized
by sperm,
i.e.
parthenogenetically. This
many
The
first
steps in the
a mass of
little cells
HOW
36
egg-cell.
This process
The
tgg.
ANIMALS DEVELOP
details
is
known
little
yolk
more yolk
NO
YOLK
SOME
YOLK
MUCH
YOLK
CLEAVAGE STAGE
EGG
Fig. 6.
BLASTULA
and larger
cells
cells
at the
at the top or
animal pole
is
collected.
birds'
37
opposed to the
The
total
cleavage of the
total cleavage, as
we have
less
yolky types.
may be equal
may be quite at
said,
it
The
smaller
cleavage
cells,
cuts
An example
of
this,
is
which they
lie
to develop in
is
quite
this
has
started to cleave,
HOW
38
ANIMALS DEVELOP
Fig. 7.
Spemann's tying-up experiment. (From Spemann.) The
upper half has cleaved several times and a nucleus has just passed
down into the bottom iialf, which has cleaved once
all
Even
the genes
seem
still
39
active, since
it is
it.
The
egg
is
cells.
mure or
less
spherical
cells,
it is
much
larger
is
not so
away from
how
this
(Fig.
6).
hollow ball of
The
cells
CHAPTER
III
all
the
endoderm
inside the
in between, a series
methods of folding, as
we shall see, but they all lead to this same result.
We might compare the development of embryos to
making toys by folding up pieces of paper but with
embryos, whatever the final shape, the folding always
starts by producing a three-layered gastrula, just as
if we are making toys we often start by making a
hat shape, and then go on to further foldings which
turn it into a boat or a frog or whatever it may be.
We shall have to describe some of the different
ways in which the gastrula is produced, both because
this is the most important process in the development of the embryo, and also as an example of how
the same process appears in a slightly different form
in different animals. In some embryos it is actually
a folding which occurs, but in others the layers
move into their right places by a streaming movement, sweeping across the surface and around the
inside of the gastrula like glaciers moving down a
mountain side. As a general rule, the lower thiC
different
is
gastrulation.
its
are selected partly because they have been particularly well investigated,
sentatives
of this
complexity. Thus
and partly
of gradually
increasing
the simple
and go on
to primitive
series
we
as typical repre-
to
womb,
in
which development
takes place.
The Formation of
The
the
sign of gastrulation
is
starfish.
The
first
is
not
much
is
yolk in
it
is
all
sorts
called
of
the
blastopore,
HOW
42
and deeper
or
ANIMALS DEVELOP
mesoderm
consists of a loose
endoderm
The
mass of
third layer
cells,
which
(Fig. 8).
rather
occurs in
fries
Fig. 8.
The
mesoderm
is
more
difficult
as has
their gastrulation
has
fact,
investigation that
actually
e
Fig. 9 a Development of the newt's egg. {a) The blastula in its membranes, (b) View
from underneath of the early gastrula showing the beginning of the blastopore.
(c) Later view of the round blastopore, {d) View from aljove of the slit-like blastopore
at the end of gastrulation. (e) Later view showing the open neural plate
embedding them
43
in
little
in dyes.
The
up
as coloured patches
is
on the
is
chosen rightly
cells
remain quite
its
gastrula-
to
it
but rather to one side (see Fig. 10, a). From the
outside, without the aid of coloured marks, all we
can
see
is
first
becomes bigger,
HOW
44
ANIMALS DEVELOP
slit
on the surface of the embryo, marking off a horseshoeshaped area, which is darker in colour than the rest.
The sides rapidly get nearer together, and the
horseshoe-shaped area becomes squeezed up to a
dumb-bell shape, and finally to a deep groove. The
dark area
is
known
it
has arrived.
It is
first
when
nearly
up.
all
^,
all
45
1
HOW
46
ANIMALS DEVELOP
bottom of the
of coloured patches.
Fig.
1.
{a), and Weissenberg's map for the lamprey [b). Side view; the large
arrows ma^k the position of the blastopore. Widely spaced oblique
lines, skin; close oblique lines, neural plate; dotted, mesoderm;
white, endoderm. The directions of cell movements are also shown
on Vogt's map.
showing
for
develop. It
usual to speak of
has arrived at
its
final
mesoderm
it
will
before
it
position as "presumptive
and so on.
We
can therefore
Vogt's
call this
map
map
a map of
newt
1 1
a,
47
which
also
the
is
to
line,
while the
way by
diving
wrong
situation
own
the presumptive
If
we
typical
mesoderm
so on.
we
can soon find out what the three layers develop into.
We find that, as was said above, the ectoderm forms
the neural plate, which turns into the brain and
HOW
48
central
ANIMALS DEVELOP
nervous system,
mesoderm develops
and
also
the
skin.
The
the
lungs, etc.
and there
is
not so
much movement
of the blastopore.
Lampreys
worked out by Vogt, Weissenberg has done experiments with the lamprey, and Wetzel has given a full
account of the gastrulation of the chicken. We need
not give much description of the development of the
lamprey, since
it
is
extremely
Fig.
11,^,
we
shows
the
map
49
with
Vogt's
map
is
immediately
obvious. It
ment.
It
is
Birds
The
chick
embryo
is
logical investigations.
The
first
studies
we know of
when
the
embryo
is
can hardly be made out without a lens or a microscope. The process is, however, particularly interesting, because it is in some ways transitional to
the conditions found in mammalian and human
embryos. Both birds and mammals are evolved from
extinct reptiles, and are therefore related to one
another, though rather distantly.
The
very large amount of yolk and undergoes the disooidal type of cleavage described in the last chapter,
little
disc or
HOW
50
ANIMALS DEVELOP
egg
which
is called the blastoderm, cells sink down and then
turn underneath and grow forward along the under
surface until the whole area is provided with a
second layer, with ectoderm above and endoderm
is laid.
below.
derm
it is
When
is
little disc,
is laid,
the blasto-
more
in this
chapter.
some of
51
cinematograph photos
of the development. The cinema photography can be
done in two ways Graper removed part of the
shell over the embryo, stained the whole embryo
with a non-poisonous dye so as to make it easier to
ous dyes
the other
is
to take
EMBRYO
WATCH
PETRI DISH
GLASS
WATER
COTTON
WOOL
NUTRITIVE
MEDIUM
Fig. 12.
of the
shell,
it,
and put
it
on
When
the egg
is
blastoderm has
fertilized in the
circle
being
consists of
of opaque
The
after
first
ectoderm-cells
collect
together into
a thickening
which appears in the posterior part of the transparent area, and grows forward till it stretches as a
HOW
52
ANIMALS DEVELOP
ECTODERM
MESODERM
ENDODERM
Fig.
13.
Section
moves
down
into
it
from each
If a
little
finely
the streak.
length.
drawn
The movement
telescoping
53
The
streak, in
up along
its
and
becomes
thinner and longer this part separates off from the
rest of the streak and becomes the neural plate and
the mesodermal rudiment of the backbone. This
mesoderm does not remain as a flat plate, but soon
breaks up on each side of the neural plate into a
series of little lumps, the body-segments, which are
found in all embryos at this stage, but are more
obvious in the chick than in the other embryos
which have been described. Between the two rows
of segments, immediately beneath the neural plate,
is a continuous strand of mesoderm, which is the
first part of the backbone to be developed.
In a
map
as
it
is
fully
the
is
HOW
54
ANIMALS DEVELOP
Fig. 14.
Map of the presumptive areas in the chick,
in the late primitive-streak stage, aged 18 hours. Only
the transparent area is shown. The arrows show the
directions of movement and the primitive streak is
drawn black. Widely spaced vertical lines, skin; close
horizontal lines, neural plate; dotted, mesoderm. The
endoderm lies underneath the whole surface
blastopore there
is
Mammals
We
must now go on
mammals, but
it
will
to consider the
development of
55
mammals
are derived
In the
first
from
yolk,
reptiles
all
the necessary
stage with
its
develop-
hypothesis, but
it
ancestors were
The
it is
its
attached
and allows
oxygen and food substances to diffuse from the
maternal blood into the embryonic blood, which is
to the mother.
placenta
is
porous,
which
is
sustained.
little
yolk
a different way.
it
arrives at
The
cleavages
HOW
56
ANIMALS DEVELOP
lump
the inner
is
cell
embryo, while
into the
all
and corresponds
out in
all
directions
till it
it,
forms in a similar
way by
differentiating
between.
It is
not
known
exactly
some mesoderm
derm
is
57
INNER CELL
MASS
ENDODERM
TROPHOBLAST
ECTODERM
Fig. 15. The development of the rabbit, {a) and {b) are sections through
the whole egg showing the formation of the endoderm, {c) is a surface
view of the primitive streak which develops on the inner cell mass, and
(d) (from Quain's Anatorny) is a much later stage showing the gill slits
HOW
58
ANIMALS DEVELOP
out such a
map
may be
embryos in cultures
like the
possible to
work
by growing the
chicken embryos.
It is
for a
technique
of cultivating
remain a useful
but
it is difficult
scientific
to
imagine
The
human
Gastrulation
is
it is
way up
59
ties
womb,
in the
as
it
mammalian embryos
quickly
till
slowing
down
the
main
parts of the
body are
man
present,
at least, for
embryos
and then
gastrulate and two
gastrulation,
to
like the
to
A human
embryo
at the primitive
in later stages,
is
takes
and speed
HOW
6o
ANIMALS DEVELOP
development within a
evolution
womb
is
mammalian embryos
a fairly
new idea
in
The
is
womb
provides.
different in different
mammals.
On
not a
strict
is
an
infant,
live
its
lives is slacker.
own apprehension
mouse
feels
Perhaps
of time, so
a minute of man-time
time.
CHAPTER
IV
first
are not
we have
nature,
how
this
is
an
them.
at
It
fertilized.
But
it
is
something simpler;
better to
let
us ask,
begin investigating
why
it
develop
may be
in
accordance
with
pre-determined
62
HOW
ANIMALS DEVELOP
altogether. It
is
fate
unlikely that
all
it is
tative pole
63
the
number of
is
The
which were
experiments
first
successful
embryo were
He
by Spemann.
carried out
cristatus,
taeniatus, lays
Spemann
in
a rather dark-
first
mm., or a tenth of an
special instruments
one
is
made
much
the other
holder,
tions
inch,
is
little
microscope.
The advance
of experimental embryology
is
always
waiting on two things, the invention of new instruments and methods, and the training of people with
sufficient
manipulative
skill
first
to use them.
Spemann's
for
HOW
64
ANIMALS DEVELOP
really
but
it
skilful
difficult to
use moderately
become
with them. For working on chick
embryos, which will be discussed on page 70 f, Spemann's instruments are useless because the tissue is
too tough to be cut with the fine glass points; one
has to use coarser steel knives, which are rather
difficult to
more
No
handle.
ments really
embryos, and that is partly why we know so little
about them. The tissue is so sticky that it clings to
the end of a knife, and if one tries to scrape it off, it
usually gets torn to bits and spoilt; this small technical difficulty makes the most important experiments
impossible to perform.
the
Embryo
is
Integrated
He
taeniatus
like its
surroundings;
65
Spqmann's
16.
Spemann).
{a)
into
66
HOW
self-differentiate
ANIMALS DEVELOP
situations.
When
the experi-
little
skin.
cristatus
finished
determiner,
then,
must have
its
the parts
The
action
to
first
experiment.
He
He
took a young
it
in
its
new
position.
glance at Vogt's
map
of the
in fact, in
its
half;
Spemann
suggested that
it is
67
actually
Mangold,
The
it
Hilde
his pupil,
cristatus
gastrula
and
presumptive
fate,
while the
part
neural
of the host.
The mesodermal
of the
tissues
was
which
It
is
strongest just
this
map
capabiHty
is
and becomes
Pr. med.
Sec. med.
sec.
^7-An organizer graft in the newt (from Durken, after Spemann and
(a) Shows the neural groove of the host embryo with the induced
neural groove on the left, (b) Shows the induced neural groove more clearly.
(c) Is a later stage, the host's neural tube is on
the right and the induced neural
tube runs straight up the middle, (d) Is section with the host's neural tube on
the left {pr. med.) and the induced neural tube {sec. med.) on the right, with an
\l^'
Mangold),
induced ear
(/. sec.
aud.) beside
it
and.
name
69
Spemann gave
it.
the
determine in
this
way, and of
organization centre to
The
organization centre
is
know,
sinks in
we
mesoderm
Finally the
inside.
it
immediately above
it
to
is
all
become neural
plate, while
The importance of
part
causes
it
One way
is
to point out
of the gastrula-ectoderm to
is
important
The
better conveyed
is
actually chose
respect to
it
which
is
all
the rest
is
it
is
organized. In order
name
for
his
discovery
and accurate.
may
is
at
first
Spemann'
sight seem
HOW
70
ANIMALS DEVELOP
Birds
This discovery
is
on
to see if
it
so
and
try to
in
we might
expect,
two groups.
We
the
first is
fact,
two bird
concerned with
of the mesoderm.
All the experiments with birds were done
on
first
experi-
no
possibility
when
of
the tgg
is
transplanting
is
laid, so there is
the
''endoderm-
e
4)
5
5
i-
"-^
.-3
QJ
O in
V o
f^
Si
bfJ5
CO
:y^m
o
(K4
<^^
^^
^..'
r^,^^^ ^^^,
ORGANIZATION
CENTRE
THE
blastopore." Instead,
when
stage
just
at a
appeared
the
is
streak
now is
HOW
72
ANIMALS DEVELOP
Mammals
Organizers have been found in other groups
We know
still
rather
the
that
mammals
mechanism
essentially
organizer
difficulties
the
same
make
is
in
also
it
grafts
of the technical
because
mentioned before
(p.
ment
also
group. Probably, as
stance,
and
this
is
Among
was described
the blastula
less
it
equal in
just
is
earlier,
The
sea-urchin's tgg^ as
more or
the bottom of
size,
and
in gastrulation
inside.
Gastrulation
THE
occurs
quite
<t^-^^
soon
attempts to
cells
is
make
after
when
fertilization,
rather large;
still
73
and
if
the
one
of
earlier,
when he could
between the
cells
cells.
cut in
experiments were
into sixteen
still
cells
of four small
embryo
and
in half in various
He
at the
bottom a
cells,
so as to produce a
normal
larva.
He
also
found that
newt.
Insects
ECTODERM
ENDODERM
75
amount of
yolk.
The
nucleus, after
The
lies
is
it
fertilized,
^'cleavages" consist of
any accompanying
a stage
is
Then
is
filled
up with
yolk.
in of the blastoderm-skin at
(Fig. 20).
and
in this
way could
that there
is
an
essential
He
found
region in the bottom end
He
called this
do
is
What
which
lies
it
does
further
centre,
lies,
in fact, just
HOW
76
ANIMALS DEVELOP
where the embryo begins to be formed by the thickening and sinking-in of the blastoderm. One might
almost say that
it lies
is
doubtful. In
any
is
is
DIFFERENTIATION
CENTRE
ACTIVATION
CENTRE
much more
like
is
em-
bryo
may be
determines
how
first
and
it
is
how
done.
at
least
The
dif-
77
wave of contraction passing through the yolky core of the egg, and as the
yolk contracts away from the outer skin of the egg,
a hollow is left in which the blastoderm cells rapidly
collect and begin to develop into the embryo. If part
ferentiation centre starts a
of the egg
is
formed
if
wander down
and
set free
through
to the
way with
the
yolk
to
the
differentiation
centre.
There
is
class apart.
all
become
active
when
when
it
is
endoderm which
first
is
is
is
is
to say,
true both
being formed, as in
when
as in the
CHAPTER
are
also
stimuli.
developed
The
as
responses
to
organizing
One example
endoderm
which induces the primitive streak, which then
becomes an organization centre itself and induces the
neural plate. Many similar examples are known. In
the newt, for instance, the neural plate closes up to
form the neural groove, and the two sides of the
groove eventually join together at the top and cut
there
is first
an organization centre
in the
79
We
embryo
to
we
power, but
find that
BRAIN
SKIN
Fig. 21.
^The
grammatic
if
we
is
something
Making
Special Parts
Once
They
are, in fact,
does not
mean
instance, has
capable of
self-differentiating.
This
its
HOW
80
ANIMALS DEVELOP
It
means that
if
lump of
the
will
develop
lump a good
deal of
all, it
removed the legrudiment will nevertheless be able to form a complete leg. A system like this, where the various parts
can be changed round or taken away without
affecting the result. If a piece
is
is
whatever
tissue
is
way and
never
is
therefore, after
some
tissue
is,
in
some way
Once, when
was working
in
Germany,
met a
who was
full
Communist
the
He
party.
arises as a small
gastrulation occurs)
is,
own
its
As soon
job
is
main
as the
some
special function,
and
own
limits
and
elect their
own
them can
self-differentiate
and
act as
out
much
centre. I
harmonious
own show
with-
to say
way
amazing how
fully and in what detail it develops. A very good
example of this is seen if the rudiment of the legbones are removed from a chicken embryo and grown
and
starts
self-differentiating,
it
is
embryos. Miss
has
made
Fell,
from this
film. When the rudiment was isolated from the
embryo, it was a little shapeless mass of mesoderm.
and the
pictures
shown
HOW
82
From
the
this
cells
ANIMALS DEVELOP
had turned
and were
we
1 1
1),
at the joints
right shape;
surface.
develop
right shape
and
bone
also occurred.
to degenerate.
83
Mosaic Eggs
The
stage
of development
described, in which
which has
is
most embryos
been
all
just
is
independent and
this stage
is
do
their work.
The
organizers cut
eyes,
ears,
some
eggs, to
is
it
is
fertilized.
sea-squirt,
egg
is
which
is
formation
They are known as organforming substances. We do not know what they are,
but in the eggs of some species of animals they have
of some particular organ.
be
is
can
easily recognized.
that
if
they are
which they
HOW
84
ANIMALS DEVELOP
abnormal
removed from
this
up
or
if
fail
to turn
at all.
is
it
is
quite
85
way
about.
come from?
very
it is,
like
an
there the
will
appear
later,
just like
As we
important things about the organizer is that it contains an active chemical substance which gives the
ectoderm the necessary stimulus to make it develop
the substance
It is
is,
We
it
all
inactive form
is
HOW
86
ANIMALS DEVELOP
The
is
tissues.
we
how
shall
this
activation happens.
The
i.e.
the deter-
shall
rate, partly
by the
The
of organ-forming substances
localization
in
Human
eggs
not
are
mosaic.
They probably
chick organizer
streak stage.
days
is
active, that
The embryo
after
conception.
is,
at the primitive
then,
separate
87
on
specialized in producing
always
many
quadruplets.
When an embryo
Structures
too complicated to be
summarized
book
only
of
as this.
how
the
There
main
is
room
in such a short
for a description
88
HOW
down and an
idea of the
mental pattern
excellent
way
called
is
books,
Embryology and
ANIMALS DEVELOP
such
forth.
as
Graham
in
MacBride's
Invertebrate
where the details of the later processes of development are described. The reader who consults these
books will be amazed at the mass of knowledge we
have about the development of the embryos in
different animal groups.
It has been said above that, as a rule, embryos of
different kinds resemble one another the more the
younger they
are.
This
is
is
them
in quite dif-
in their
Now
all
light,
known
as
89
And we
find, if we
study
way
(Fig. 23).
The whole
sinks
retina
inwards and
IRIS
^^ '^
SKIN
LENS
RETINA
of skin grow up
all
The
though
it
become con-
HOW
go
ANIMALS DEVELOP
Larvae
In
many
is
The
best
known
tgg develops
becomes
is
found
Everybody knows
that a
illustration of this
it
first
into a caterpillar
and
much
Some
insects
have
little
is
still
a few
extremely
up
to
discs
is
91
very complicated.
Fig. 24.
Part of a caterpillar v/ith the skin dissected away
to show the buds from which the butterfly's wings will arise
have a water-living larval stage, the tadpole. The Ascidians, which we mentioned before as
examples of animals with mosaic eggs, are not so
well known as frogs, but they also have a larval stage,
which is rather like a tadpole. Here the metathey
still
HOW
92
aquatic.
The
ANIMALS DEVELOP
swim about
tadpoles
freely,
but the
we did
not
know what
like.
why
the life-histories of
some animals should be complicated by the development of larvae forms when the animals are finally
going to turn round and become something absolutely different. To develop into a butterfly by first
becoming a caterpillar is rather like going to Birmingham by way of Beachy Head, as Chesterton has
it. Probably it serves a useful purpose by making the
embryo able to support itself before its development
is complete, so that there is no need to store so much
yolk in the egg to keep the embryo alive. But to show
efficient or perish.
so
it
more
would
force the
specialized
of the adults,
embryos or larvae
and complicated.
to
become
%l
Fig. 25.
CHAPTER
VI
the
Organizer Works
is
only a
plate.
It
stimulated
it
ready to do
so.
The
when they
are
We
this
is,
they
HOW
94
cess
ANIMALS DEVELOP
petencies
and
it
by forming
other organs.
Ahhough we know
what makes a
tissue reactive,
we have found
out
first
still
effective
it
When
material
form an
im-
But
way from
organizers.
those
The
of an embryo,
or a
it
is
tail
it
which
are
induced
by
lining
is
part
i8).
The
all
the different
95
to be the case. It
neural plate
shall be.
it
The chemical
stimulus
is
which
determines which part of the neural plate shall be
induced
is
else
The
fact that
covery merely
why some
us
tells
particular part of
the
is
with respect to
is
it
this process,
in a competent
it is
acted on by the
embryo because
the whole
it
within
is
The
the organizer
is
active as a determiner,
of the individuation
which it is active
the embryo.
field
and
the
embryo together
complete animal.
which
emphasises the
way
in
General Pattern
parts of the
in
Particular Causes
It is
way
It
to
fits
the different
make up one
single
HOW
96
ANIMALS DEVELOP
determines where
the
arise,
and then
where the
We
its
It therefore
and
how
so on.
effect,
shall
We
eye-cup
is
effect
is
is.
if
we add any
will
do
its
best, so to speak,
leading.
Hump
which you
down
the
Hump. The
in the picture,
is
sidings.
to a set
first
point,
two trucks
The
organizers,
last
chapter,
and they again sort out the neural plate trucks into
brain trucks and spinal column trucks, and the
on through
all
97
trucks.
And
the organizers,
till
we may
so
all
made up
this
model a competent
tissue
is
analogous to a
is
it
runs
down
this siding
way
or the other, so
it
as
pare
with
is
individuators
the
means determining what detailed structure the induced organ will have, it means settling whether it
be brain or spinal column, etc. In our model,
therefore, it means sorting a whole set of trucks into
their final sidings so that each siding gets the right
number. The first point, or primary organization
centre, can only do that if it is connected in some
will
way
by a system of
levers so
of individuating levers.
The important
is
thing to
remember
in this analogy
happen
one
one
single
truck, that
small piece of
tissue.
to
is
to say, to
HOW
gS
ANIMALS DEVELOP
tion
is
combine
to
The
essence of individua-
recognize
it
is
induced, whether
we
column,
we know
of always
fit
of a brain
fitted
We
harmoniously as possible.
is
make a complete
of the individuation
by a man
field, as
It is
99
in the
developmental
first
how
soul.
it is
is
stand
the
not understand
all
about development
yet,
they
still
same
The
the
set
of trains to be
made up
raw
to build
The
up
on
different days.
individuate.
Some of the
we know of seem
is
so
Seidel's
HOW
100
ANIMALS DEVELOP
does individuate.
Regeneration
In
many
through
animal
is
animals
life.
Their
effect
badly wounded,
is
fields
seen
persist
all
when such an
example.
and builds up
again by regenerating the part which
a complete leg
has been
individuation
In
lost.
this
to those con-
For instance,
cap of
if
cells called
an adult newt
loses
a leg, a
little
on the
field
it
develops out of
legs
is
it.
It is
if
necessary. If
it
part of the
now
tail
tail
individuation
In
many
all
the tissues
seem
to be
lOI
develop on
its
own
into a
phenomenon of reproduction
facts
The
when any
part of the
gradient
is
isolated
formity with
itself.
Axial Gradients,
first
drawn
to their
of,
CHAPTER
VII
all its
of an animal
is
not finished as
The
details
of
and adjusted
to one another. Organs like the heart, which are
required early in embryonic life, get a start on
other less necessary parts and are disproportionately
large in the young embryo. The control system of
its
structure
must
still
be
fitted together
may
demands made
How
The embryo
Embryos
are
The
Fed
own
its
it is
nourishment.
is
digested
and then
growing animal, is a
very complicated one. Until recently the whole subject was a chaos of unrelated facts, but a few years
built
ago
up
Needham brought
together
pieces of information in a
work
all
the scattered
entitled
Chemical
this
IO3
is
different
the dif-
live
make
by
satisfied
salts,
fuel,
(4)
first
three.
oxygen,
(2)
water,
(6)
The
for a developing
proteins,
**flesh-forming" substances.
the
(i)
real
The
enough
not contain
it
by creatures which
provide sufficient salts in the egg. But the most
difficult evolutionary step, from the embryological
as from every other point of view, must have been
the conquest of dry land. Here there is no useful
substance, except oxygen, which can be absorbed
from outside, and the embryo has to be provided
not only with its organic materials and salts but also
with its water. Amphibia, which are the most
therefore can only be inhabited
HOW
104
ANIMALS DEVELOP
up.
The
drought problem
solves the
only workable
filled.
if several
fairly satisfactorily, is
carbon-dioxide, which
to diffuse out
of waste
is
One
like all
other
waste product
is
through the
shell.
The
breakdown of
this into
itself.
is
it
proteins. Water-living
ammonia which
embryos turn
What
they actually do
IO5
is
to convert
as urea, or, in
acid,
which
aric
is
shell
is
it.
this
excreted as ammonia, as
it is
the nitrogen
in fish, then
it is
is
con-
and
is
become
modified to produce less nitrogenous waste they do
this by using more fat and carbohydrate and less
protein, as fuel, and they get the added advantage
from this that the fat and carbohydrate give rise to
some water when burned and thus help with the
uric
acid.
Birds
reptiles
have
also
So
the
far
special
in uterine
is
way of
life.
In a sense, the
The
HOW
I06
ANIMALS DEVELOP
instance,
it is
for
only in the
mammals
Even
in
its
that
is
at all
never
through the embryonic bloodvessels. If it did, not only might the mother's female
sex hormones disturb the development of the sex of
circulates actually
it,
types of placenta
passage of substances
pearance of some or
is
made
easier
mammals
the
by the disap-
107
Fig. 26.
embryo
and
in mice, rabbits,
figure. {A)
maternal blood
lies
down and
the
and there are only three layers left separating it from the embryonic blood. In the same series
of animals the wall of the placenta becomes more
placenta,
HOW
I08
ANIMALS DEVELOP
which
man
branching
the
fingers, con-
taining blood-vessels,
maternal blood
left
blood-vessels.
passes into
blood
its
is
not
which the
embryo requires for its growing organs. At the same
time the placenta passes back to the mother the
waste products of which the embryo has to rid itself;
in the more elaborate types of placenta this backward passage is so efficient that the embryo never
develops a functional embryonic kidney, which other
embryos have evolved to deal with their excretion
again into
the
substances
particular
Not quite
all
mother's blood.
uterine milk,
uterus,
chiefly
is
ready.
Some
which
in
is
is
by glands
in the
the
wall
of
the
uterus,
the
connective
IO9
as to prepare a space in
in
its
mother's
flesh.
which
to
develop embedded
start
on
this
it
is
between the
embryo and its mother is so highly developed and
specialized in Man that no further evolution along
these lines seems conceivable.
It is often the failure of such a specialized organ to
be able to evolve to meet new conditions which leads
fertilized.
In
fact,
the
connection
suggested that
Man may
Growth
Many
life
long. It
is
man becomes
we
say that
is
And
often no point at
over. This
is
which go
and therefore
size,
life
is
and
no
HOW
gets slower
ANIMALS DEVELOP
and slower
that in man the upper part of the body retains more nearly the foetal
proportions, whereas in the gorilla the legs remain more unchanged
while the head, and particularly the brain-case, becomes very small.
(After Schultz.)
how one
we know from
fossils,
Relative Growth.
The
either so as to
or in response to the
to do.
An
alteration
a small lens
lens
grows
is
fit
in with
its
III
surroundings,
due
is
seen if
faster
than
it
make
up a harmonious but middle-sized organ. The influence of the second factor, the amount of work
required, is shown by a comparison of tadpoles
large eye
more
The
Influence
of Function
to
it
its
HOW
112
ANIMALS DEVELOP
Fig. 28.
Differences in the guts of tadpoles fed (a) mainly
on vegetables, (b) exclusively on meat. (From Diirken.)
thing
is
is
stretched,
any minute
particles within
it
Now
it
embryo are sometimes filled with a jelly-like substance which is made up of little elongated particles
which get orientated
in
various
up
down
Many
movement of
to
cells will
of orientated particles. In
lay
II3
often
this
way
move along
lines
the tensions
may
arrive.
The nervous
system, which
is
be
it
move
The developing
are fixed
is
more
rigidly fixed.
is
^^^
114
Once
ANIMALS DEVELOP
the
altogether,
fibres
On
the organ
thus
it is
an organ
In view of this
is
cut.
it is
although well
Hormones
Much
It
is
Fig. 29.
Effects of the pituitary gland, (a) Giant
with over-developed pituitary. (From Engelbach.)
{b) Dwarf with under-developed pituitary. (From
Behrens and Barr.)
Fig. 30.
those of
suffering
from hormone
diseases.
(From Stockard.)
we
shall see,
some of these
effects
I 1
can be brought
about by other non-specific substances. The hormones are usually manufactured in special organs
body
which are
called the ductless glands, because they have no
opening to the exterior of the body but pour out
set aside in the
The
make
still
be found.
is
One
of the most
is
on growth.
If
it is
in
animals
HOW
Il6
ANIMALS DEVELOP
it.
evocator-substances
It
I17
seems as
if
they
is
is
such an important
ele-
ment
in our
elaborated,
whole constitution,
either male or female. These two phases
making
it
its
the
fact
about the
among
other things,
fertilized
and
emphasized in
One
sex-cells
is
that
new development
fertilized
egg produces,
so
a continuous succession of germcells, the later ones derived from the earlier ones by
division and reunion in fertilization, so that they are
is
same
may
an immortal piece
HOW
Il8
One can
only
ANIMALS DEVELOP
make a
is
Now
it.
in
many
cases
may be
a particular
is
set
is,
in fact,
cells
in the
organizer.
In
body, that
this
sort
is,
in
response to some
young embryonic
cells are,
who
wishes to can
now make
II9
an actual material
living thing by endowing a staff of experts to keep
mortaUty of part of himself
Each
as
it is
offers.
cultures
little
few hundredths
is
its
than 6 months.
We must now turn to the second question; what
decides whether the germ-cells, and the whole
animal, shall be male or female? Actually two groups
less
One
set
ment.
The
hereditary
which
its
basis
influence
arise later.
probably always
is
is
overcome by the
The important
thing to
is
maleness and femaleness, sometimes with one preponderating, sometimes with the other.
tary factors or genes for maleness
The
heredi-
carried
HOW
I20
ANIMALS DEVELOP
that a special
sex factors.
of two
X to
Xs
The
have one
will half of
too
them contain an
many Xs
may be
too few
"super-females,"
Xs when we
when they
or again
get
"super-
121
and finally we may get proportions intermediate between one X and two Xs to the set of As
and then intermediate forms or ''intersexes" develop.
Again, there are strong and weak varieties of the
male and female factors, and unbalanced intermediate animals can result from some combinations
of them. Most of these abnormalities have been
found in insects, though they may perhaps occur in
other groups where they have not yet been discovered. They are nearly all infertile and generally
males"
feeble.
The
arise
the matter at
discuss
to go
more deeply
fully.
into the
all
The
Genetics of Sexuality in
122
The
Development
Activity of Genes in
we have got an idea of how development comes about, we must ^o back to the question
Finally,
now
that
development?
emphasise
No
is
the
How
2.
do the hereditary
The
enormous importance of
genes,
its
father,
into.
which
determine
Which
realised as
to
point to
their effects.
first
inherited from
it
all
The
it
nature
its
mother and
it
may develop
en-
Human
personality
in affecting characters
and
by environment in upbring-
in-r
man
develops
five fingers
by the environment.
It
or four
is
very
but whether
little
affected
anatomical abnormality of
this
is
stages of
development
123
a relatively
stances.
may be
It
we think only
as the
of a single lifetime
is
gradient of yolk.
portant regions
are
The same
is
we mentioned
in insect eggs.
But there
Firstly, these
when
when the
is
to say,
page
77).
Secondly,
is
Now
during
this
period of
same
as the
cell in the
mother's
body.
is
egg matura-
formation of imperfect
HOW ANIMALS
124
in question to be sterile.
DEVELOP
way
examples
is
One
of the most
famous
which some
and others
in a right-handed. This difference depends on the type
of egg which a given mother produces. That in its
turn is determined by the genes which she contains.
individuals are coiled in a left-handed screw
We
then,
see,
itself
is
It is interesting to
when an
petent
cells
which have a
cells
which react
an organiser
to
their
own
is,
in
achieve
is
to select
this
range of
possibilities.
What
is it
we need
to decide
pens in development.
rials
by
has led us
far the
first
Now
of all
them such
To
answer
cells are
the
which go
125
There
are, of course,
many
make
it
seem reasonable
to
compounds
fats and oils of
other classes of
(i.e.
suppose that
all
these sub-
working hy-
as a
is
the production of
which may
act as
In the
last
on trying
trated
to
which genes are able to control the formation of parVery great advances have been made
and we can now say that we have at least an
ticular proteins.
in this,
to
it
is
essen-
has not
do not seem
bryonic development.
instance, have
lines
which
Some major
come from
em-
contributions, for
crystal-
126
modern
technical
germ
cell, as
we
beginning of chromosome
to the
now become
has
1,
and
known
is
joined
vice versa.
as
recombina-
in genetics. It
is
if
two genes
on a
lie
some
they
if
lie
is
much
less.
We
stretch of
chromo-
how
how often
recombination happens between them. One of
far
the
the
study recombinations
rarely. Bacteria
when
it
became
possible
highly elaborate
techniques,
it
many
became
127
which were
and rejoining
once in a million.
If
we can
two
effects,
lying so near
fact,
is
made
is
about that of
The
sort of chemical
compounds
is
to
determine what
of
essential for so
thought at
much
first
would play
that they
cell, it
might be
the essential
is
actually
been possible
from one
to
strain to another
it
chromosome
is
It
it is
the
in determining
has
it
on
its
made up of
hereditary
effect.
HOW ANIMALS
128
end
to
DEVELOP
end
to
form
The
a chain.
evidence about this comes largely from X-ray crystallography. This shows that the type of nucleic acid
found
in
chromosomes (known
DNA
acid, or
as
deoxyribose nucleic
more
It
con-
sists
as a thread,
each other in a
spiral.
at a particular place in
if,
is
given constituent molecule or nucleotide, this immediately determines the nature of the nucleotide at the
DNA, which we
up from
pairs of nucleotides.
This
fits
nation which
The
we
from recombi-
stretches of
DNA
which the
size as
the smallest
two
The
ent picture.
Now we
of
the
DNA
cell.
nucleic
have to go on to consider
how
this
thread
rest of
The
acids,
first
point
are
essentially
is
like
molecules.
built
to
129
are attached
known
as
amino
acids.
It
DNA
in the
at the proteins
One
is
case in
that of
which
haemo-
known
genes are
man which
in
of these
produce abnormal
abnormal haemoglobins
differ
nucleotides,
tides
of
an alteration of one ol
first
place, to
and
this altered
sequence of nucleo-
amino
We
acids.
still
nucleotides in
DNA determines
acids in protein. It
is
this
sequence of
the sequence of
DNA
is
is
amino
not a
normally
if it
it is
130
On
this
One can
an amino acid
into the
medium surrounding
a cell which
is
engaged
is
being produced.
embryonic
cells
It
is
sites at
which the
the
major
site
of protein
are engaged in
making
the
it.
main
which
cell,
the labelled
first
amino
in the cytoplasm.
The
actual
sisting of particles of
size. It
turns
amino acid
consists of a
group of very
known
as
microsomal
131
particles.
These
They
of
It is clear that, if
it
passes
from gene
to
is
is
tein.
is
also
of the hereditary
The
DNA
nucleotide building-blocks
contain a sugar
known
as
The nucleic acid in which this is incorknown as RNA for short. It seems very
sugar ribose.
porated
is
somal particle
is
also seems to
actually a kind of
RNA,
although
RNA
it
play
know
be seen from
this
of course,
still
many
details to
be
filled in,
even in the
132
HOW ANIMALS
simple story of
DEVELOP
influences
as yet
made
is
correspond-
is
problem, which
its
more
difficult
The
less progress.
and
in
still
is
fields of
another
which we have
developing
cell
not
is
On
on in
is
the contrary,
muscle
it
must be
produces, in
cell
fairly
definite
amounts
very
idea
of synthesis
that the
a functioning muscle
quantitative
is
how
it
there
cell,
relation
little
perhaps
up
is
between
the
We
have
contains.
at present
it
is
little
more than
a guess is
electron microscope
is
with
this matter. It
is
that
we can examine
now
much
to
at all. If
we suppose
some way
to
I33
Summing-up
This book started by considering the
classical prob-
We saw how
the
mere
descrip-
to the reca-
pitulation hypothesis
of the funda-
mental pattern of
We
how
in outline
then saw
the
arise.
on in the egg which produce organ forming substances, or evocators and competent tissues
Processes go
which react
to
give
which
is
aspects.
One
is
the "substance
com-
cells are
The
and organ
other
of the
cells of
is
body has a
tural organisation.
The
definite shape
Each
and
cell
struc-
progress of understanding in
main
a digging
down
into
We
last
chapter
how
how
far
we have
hereditary genes
as
The
per-
But
134
We
must
on whatever
we
see
around
us.
We cannot claim
to understand
how
how
the various
comes
to
cult
that they
related.
still
At any
is
how
we have
We know
to rediffi-
something
increasing
all
the time,
all.
rate,
about
edge
member
how
know it
about everything how
we
all
shall never
67
185
INDEX
Activation
100
centre,
75,
Gastrula, 23
gene, 32, 119, 122
germ cells, 24, 117
organism, 14, 15
orientation of particles,
animal pole, 27
germ
germ
armadillo, 87
gill slits,
ascidian, 83, 91
axial gradients, loi
growth. 109
103
layers,
track,
24
1
112
20, 57
Harmonious
equipo-
80
Biogenetic law, 20
tential system,
hereditary factors,
119, 122
hormones, 114
blastoderm, 50
blastopore, 23
blastula, 23
budding, loi
butterfly, 109
Communist
party, 81
competence, 93, 100,
123
cultivation, 51, 58, 118
cytoplasm, 27
Determination, 62
developmental fate, 61
Immortality
cells, 117
of
Parthenogenesis, 35
phosphatase, 82
placenta, 55, 106
germ
48,53
primitive gut, 23, 41,
46, 69
primitive streak, 52, 55
protoplasm, 16
immortality, piecemeal,
Recapitulation, 20
recapitulation,chcmical,
individuation, 95
induction, 67
insects, 73, 90, 100, 109,
105
reduction division, 30
regeneration, 100
reptile, 49, 55, 104
116
Joints, 87, III
Kidney, 82
self-differentiation,
79
Lamprey, 48
larva, 90,
leg,
1 1
80
lens,
79
86
of development,
59
sperm, 26, 62
localization,
sp>eed
Mammals,
man,
Ectoderm, 24
egg, 26
cndoderm, 24
entelechy, 98
microsurgery, 63
Tadpoles, 1
turtle, 104
monstrosities, 59
mosaic eggs, 83
twins, 37, 86
Fertilization, 32
fish,
mendelism, 34
mesoderm, 24
mutation, 32, 39
frogs, 42, 91
function,
1 1
51
1
tying-up experiment, 37
49
food, 102
63
sorting yard, 96
differentiation centre, 75
ductless glands, 1 15
62,
Octopus, 88
organ - forming
sub stances, 83, 118
Vegetative pole, 27
virgin birth, 35
Wasps, 87
waste products, 104
Yolk, 16
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How Animals
Develop
scientific
modern science
its kind,
animals
develop
embryonic differentiation." f.
''Mr.
embryology.
of experimental
h.
a.
underlie
the
way
in
which
processes of
Marshall, Nature
though necessarily simplified, account of his subject ... For the student
book
this
will
"If
tific
will
provide
some fascinated
in
hours.
And
it
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