Download by Booij (Leyden). Introductory § i. systems § 2

The protoplasmic
a
regarded as
Membrane
Complex
System
by
H.L.
Booij
(Leyden).
CONTENTS
Page
Chapter I.
Introductory
§
i.
Colloid systems
§
2.
The
2
of the
structure
§ 3.
Antagonism of the ions
§ 4.
The action of
§ 5.
Permeability
§
Can
6.
the
complex
§
7.
§
§
1.
2.
II.
...
coacervates
on
.
.
be
regarded
as
a
20
the
to
experimental part of this research.
with
Experiments
Lathyrus
23
pollen
Introductory
26
the germination
§ 3.
The influence
§ 4.
The influence of
§ 5.
Stability
of
of the
27
salts
germination....
29
organic substances upon germination
33
some
on
grains
37
§ 6. Respiration
Chapter III.
The
42
influence
bakers’
§
§
1.
2.
of
rate
of
influence of salts
§ 4.
The
influence of
§ 5.
The
volume of yeast cells
§
Respiration
Chapter IV.
§
1.
2.
§ 3.
on
fermentation
The
§
compounds
43
the
§ 3.
6.
various
yeast
Introductory
Measuring
organic
compounds
in
57
salt solutions
....
of yeast
57
59
the
experiments
be
from
can
The
protoplasmic
membrane
protoplasmic
membrane
Summary
55
General conclusion
Selecting
Literature
44
fermentation
on
of the membrane
The
13
16
membrane
system?
Measuring
5
10
organic substances
protoplasmic
Introduction
Chapter
membrane
protoplasmic
which
the
structure
derived
of
60
Lathyrus
of bakers’
pollen.
yeast
.
...
63
66
71
75
CHAPTER I.
INTRODUCTORY.
§
Colloid
1.
ample
systems.
fact
The very
of
modern
the
for
justification
those
only
medium of which is
their electric
sol will
sol,
now
we
sooner
now
of time,
In the
a
what
have the sol
shall find the
of
work
similar
stability,
particle
diffuse
note:
and
b.
a
water-shell
is
con-
often
oppositely
secreted.
u,
12,
are
the
into
as
The high,
relative
proof of the existence
same
are
not
direction.
hydrophobe sols;
visible
we
to
13). If
mixes
one
liquid layer,
factors
two
shall
then
electrolytes
hydrophile sols,
very rich
which
under
Dehydrating
ultramicroscope.
two
on
vicinity
causing the presence
greater sensitiveness
a
directed
the immediate
under the
charged,
There
the ionized and other
are
particle.
accepted
in
viscosity,
(to,
the
second factor
as
that the micelles
points
visibility
such
a
water
from it, thus
around
hydrophile
convert
well-defined
be
factor.
lower
at a
of
greater in
away
Also, the fact
low relative
are
act
given lapse
vital
a
higher, and the univalent
dipoles
surface,
influence is
than further
Coacervates
which
may
can
a
the
at
The
hydration.
ultramicroscope,
media
act
be
charge, hydrophile sols possess
of these sols
hydration.
the
to
electrolytes
a
the highest concentration. In the case of positive
valency rule will be found for the anions.
This
of
to
such
at
called
groups.
a
minute,
stability is due
concentrations of various
negative sols the trivalent cations
polar
viscosity
biological
of
water
Their
valency of the ions
of the micelle
of
in
suspensions
completely precipitated in
definite parts
of the
a
the
importance,
later.
or
Besides the electric
of
From
is
which
disperse
distinguish five forms of these
can
are
centration, the bivalent cations
sols,
potentialities,
biology.
of
are
particles (to —too //).
ascertain
to
we
case
cations
to
of colloids
system
charge; they repel each other and, consequently,
required
are
a
not
precipitate. Neutral salts will decrease the charge; the
having turned into a suspension of uncharged particles, will
precipitate
If
We
sols
Hydrophobe
electrically charged
a
affords
systems
water.
being
review of the
(43):
systems
Sols.
a.
closer
colloid-chemistry
standpoint,
watery
protoplasm
a
play
a
in colloids,
prominent
3
part in these systems,
caused
to
viz.
by hydration.
). When the
1
X
C
the electric
be looked upon
will
the
as
I
+
i
h
=
d
x
this formula
In
h) its
i
2
D)
of
sols, that
of
of the
sum
dielectric
complex
felt
(21).
is
its
at
of
will contain the smallest
be
present
For
charged.
in
quantity of
the
excess,
form
positive
again
can
be
the anion,
at;
a
3
—1
great
a
ions
>
following
2 —1
>
influence
Those, possessing
possessing
Thus,
we
we
a
two
1 —1
low
than
makes
itself
the numbers
of
coacervate
uncharged. Should
then the
negative
coacervate
component is
the
upon
the
cations of
should like
to
distinguish complex
as
coacervate
for anions
a
high valency work in
a
valency.
If
to
phenomena; 20).
make the
can
well
drop
negative component,
excess
as
droplets.
large
we
represent
of the cation and
a
valency
a
lower
salt
by
b) that of
parallel rules of valency will be arrived
and
1) In other colloid systems
Generally,
mixture
a
form many smaller
one
—b, a) being the valency
the
This
water.
point the
itself from
droplets with the
concentration than ions of
symbol
viscosity
this
the
seen to
frees
27). Both for cations
rule will be found;
liquid
a
much lower
surrounding medium (disintegration
21,
as
droplets in the electric field will
Under the influence of neutral salts,
disappear (19,
The
constant.
of
and
interval
positive electrode. On the side of the negative
component
coacervate
present in the
the
be
secreted
when
excess,
when
coacervate
electrode, the larger drops
the
find the
to
and be
water
in
particles,
particularly in the domain
At
equal.
example,
be attracted toward the
Here,
of the
the medium,
we
are
minimum
of the components be present
will
one
of
of the “bound”
maximum
are
' I00)
(
Hence
It is
that this
coacervation
Coacervation
n
/
oppositely charged, is
are
positive and negative charges
one
Co
(h, + h2)
constant
rest
viscosities.
partial
the
to
=
supported by the fact that the viscosity
hydrophile
the
X
Cl
d
or
layer together with the
is
due
in
i
h2
between the
particles in a given coacervate
particles, having lost part of their water-shell,
conception
refer
to
equilibrium,
Formally, this repulsion may
d) the interval between the particles.
coacervate
repulsion,
units.
two
D
the
in
equal the repulsion
e) represents the charge
hydration,
is
coacervate
particles.
of
sum
Cl
D
the
should like
we
c
attraction
tendencies of the
hydration
both factors
Jointly,
complex relations
as
the electric attraction and
1
—3 >
charge
1 —2
of
>
1 —1.
complex
Salts
also have
coacervate
drops.
high valency, will diminish chiefly the
we
may
come
across
these complex relations.
designate these systems
coacervates,
as
complex
systems.
complex films, complex gels,
etc.
4
negative charge, and therefore, they will render the drops positive.
On
the other
find the
continuous
i
replace
may
high
—
2
>
with anions
i
of the
one
—
>
i
i —2
complex
is
phenomenon
of
high valency
Tri-complex
i
>
—3.
coacervates
also
are
colloid
amphoteric
Of the
examples, known
anion
ion, the
known
densed system
gether,
ion of
an
autocomplex-coacerIt
(29).
crystal
which the micelles
can
or
be
can
a
particular
the film
group,
biology. For different
ion.
liquid,
a
con-
aggregated
to-
in
definite pattern.
a
bi-refringence of the system.
membrane is
or
which will
reasons,
as
condensed system,
a
aggregated in
are
often be ascertained from the
this
as
crystalloid
a
colloid
a
fairly closely
defined
molecules
or
coacervate
are
three
upon
depend, namely:
day, the cation is
regarding
that
is
anion.
—
crystalloid ion
a
with micelles
colloid
a
By
cation
—
this
to
up
being either
Colloid crystals.
In
by
components
called
components that the establishment of these systems
This
we
(12).
vation
c)
are
drops negative. Here,
rule:
valency
>
i
—
which
valency,
salts
action that makes the
an
following
3
We
neutral
hand,
distinguished by
be
important
to
considered later, it is
probable that the protoplasmic membrane is partly formed of lipoids
(lecithin and the like). In the
that
of
oleate-
and
electric attraction
forces
that
the
which
and
tendency
Our
which
“dehydration”,
alcohols,
and
London-Van
of
cholesterin
is
the
like,
der
Waals
which
by
electrolytes
regarded
factors:
lipophile
as
a
be
shown
molecules
It is
the
(14,
non-electrolytes
affecting
by
interpreted
force.
attraction,
Waals
der
presumption
line themselves
up,
Thus,
action upon monomolecular lecithin
determines
transition
the electric
oleic
as
obvious
acid,
an
filling,
as
it
is the
to
higher
of the
increase
that it
permeability,
(65). This
the
amount
water,
were,
of
a
the pores
15). The influence exerted by
on
a
complex
one or more
hydration
system
of the three
and
can
be
influencing
the attraction
of the
groups.
d) Flocks.
ter,
and
to
as
besides
of “sensitizers”.
means
lecithin film; the cholesterin molecules
between the lecithin
tend
the
the addition of cholesterin
likewise
may
is
well
as
powers
the London-Van
chains
be stimulated
cholesterin has a “dehydrating”
layers; the surface decreases on
complexes,
other
starting-point
fatty acid
hydrophobe
can
of lecithin
hydration, i.e.
play (14).
into
come
case
stearate-coacervates,
These are condensed systems of an amorphous characthough it is possible that they consist of very fine colloid crystals.
5
If
add
we
sensitizer
a
to
an
flocks
oleate-coacervate,
As the concentration of the sensitizer increases,
non-refringing
from the
(85).
stage
in order
Besides
The
form
to
the
as
a
whether
the
of
phenomena
The
in
of
structure
The existence
to
the
are
going
in the
on
be
or in a
liquid state. The
explained from the action
complex relations.
membrane.
membrane around the proto-
permeability,
the cell’s
is no longer a point
deny the existence of this separate
protoplasm itself.
by the experiments
to
colloido-chemical
In this
of Bigwood
theory they
who
(3),
are
that
proved
of the gelatine gel for Ca- and Cl-ions depends
charge of the gel. It is noteworthy that this permeability
follows the Donnan rule.
membrane
be
cannot
Other authors
grounds
of
are
both
Finally,
of
from
the
accepting the existence of
mation
of
This
a
the surface.
a
it is
Like
point
the
hairs
to
The
The Brownian
of the
concentrate
may
even
Most
layer is
convincing
furnished
differ
well-known
some
on
surface.
the surfaces,
the
lead
rest
to
of the
the forwith
experiments
forming
uninjured cell, parts of the protoplasm of
When the
proto-
and the refraction in the surface
from
those in the inner
proof of the existence of
by
the
properties
the
on
in the direction of a membrane
are
movement
protoplasm
contain
to
formed
impermeable to dyes.
plasm is “killed”, it gets rapidly stained.
root
on
membrane inseparable
differences with
phenomenon
membrane.
Nageli
a
endowed with other
tendency
the
membrane
appear
membrane is
explaining
surface
solid
Hydrocharis by
Hydrocharis
the
lipoids have
concentration
controversies
though
a
permeability.
to
existence of
accept the
protoplasm,
protoplasm.
relative
these
than the latter, because
Proteins and
A second fact is that the existence of this
proved morphologically.
experiments
elements of truth. We
on
a
to
permeability
upon the
this
be
is
important than
more
attributing the changes in permeability
supported
the
is
protoplasmic
non-existence of
or
control
solid
to
upon the
of contention. Those authors who
layer,
a
permeability will have
organic substances
plasm, said
reactions
rigid.
grow
complex gels
membrane
protoplasmic
complex system of biocolloids,
question whether it subsists
2.
system will
(22).
of salts and
§
the colloid
mesh-work,
a
question
regarded
change
powerful bi-refringing
complex flocks and complex colloid crystals,
known
are
comparatively
a
This
the increasing orientation of the molecules.
proves
Whenever the micelles of a sol are seen to adhere locally
Gels.
e)
to
be detected.
can
these flocks
a
Chambers and Reznikoff
layer
layer.
protective
(35,
36).
surface
6
On
Amoeba into
introducing
membrane
touched with
for
cause
KC1.
or
the
On
the other
protoplasm
hand,
the
toxic outside the cell than inside it,
are more
Amoeba
and
2
CaCl 2
find the
we
reverse
is
distilled
in
placed
Also, when the organism is placed in
NaCl
the
water,
KC1
or
is
establishment
soon
a
solution of
solutions.
According
these
to
function of the internal
a
surmise
correct,
we
that
great
the
are
not
or
2
protoplasm,
CaCl2 ,
take
place
this
re-
dependent
surrounding medium.
surface
to
permeability.
be
phenomenon
substances
capillary-active
essential part in it. There
an
the
—
a
that
assume
playing
be considered
to
membrane is
safely
can
(especially lipoids)
things
quickly
authors,
of the membrane is very essential
structure
If the
will
MgCL
re-establish itself, which does
the collaboration between the latter and the
The
former
itself.
the membrane will
on
MgCl
Hence, while NaCl and
solidify.
to
MgCl 2 and CaCl2 . When wilfully injuring the membrane whilst
re-establish
in
destroyed, when it is
2
solutions of NaCl
KC1
they found the
solution,
completely
glass needle. MgCl 2 and CaCl have no appreciable
protoplasm becomes slightly more liquid on injecting
a
influence. The
injections
NaCl
a
The latter is
soften.
to
slight permeability for
are
other
and the
water
velocity of permeation of substances that dissolve in lipoids
from
which
the
From
inferred.
deduct that
character of the
hydrophobe
this
of the
conception
composition
the
of the
membrane,
protoplasm
we
again
may
greatly influences
(69)
that of the membrane. Here, it should be noted that Lepeschkin
protoplasm
sees
the
fact
until
the
From
injured
as
siderable
that
the
a
molecules
makes
hypothesis
a
the molecules
of
the
by
will burst if the volume be
find
a
layers
It
goes
lipoid molecules,
must
too
by
(42).
tension is
believed
adsorbed
on
to
low
be caused
the lipoid
their
having
layer will possess
The
layer.
get
con-
the
(The acceptance
a
by
follow
that this
the
a
briskly.) In this
polar
film of
way
the thickness
of the
may
re-
of
rate
of volume; the membrane
of
a
groups directed
high surface tension,
value
We
in amongst the
saying
closely
expansion
expanded
ward. A similar
protoplasm
not
With the extension
creep
without
model of the membrane which has
of
does
point of
conclude that
thickness.
dissolved hitherto
the
a
possible the acceptance of another
membrane
surface extension, caused
(40, 41)
monomolecular film.
of the membrane.
establishment
Davson
multimolecular
theory: the existence of
of surface,
membrane
protoplasmic
swelling of the cell has reached
have
must
of the surface
compound of proteins and lipoids.
Danielli and
bulk,
membrane
a
—
membrane may be
not
protoplasmic
they
few
out-
shown
surface
protein-like substance,
picture
a
similar
layer
as
7
existing
the internal
on
surface. Danielli
the influence of Na and Ca salts
and Davson put
permeability,
on
of those of the Na and Ca salts that have anions
phosphate group.
or
salts
are
able
to
As
rule the Na salts
a
a
the Ca salts will
NaCl solution,
Fig.
ability than
be
not
a
membrane.' This
Ca
which
are
We, for
do
us,
tioning of the cations
of
the
first
be
to
treats
permeation
1.
the
3.
a
as
mem-
perme-
simple
a
at
a
are
rate
regarding
borne
out
by
interpretation of the func-
all that and this is
as
Various
quicker
their views
high valency,
different
permeability
high valency.
permeability
on
a
forms
dia-permeation,
2.
vide
certainly
§ 3.)
physico-chemical angle.
of
permeation
dis-
are
in-permeation and 3.
we
are
ex-
chiefly concerned
latter
The
no
2.
different groups
homogeneous
wall
permeating
of membranes
substance
the
through
does
not
let
(Fig.
2).
the solute
pass,
capillaries.
homogeneous wall substance lets through the solute; there
evidence
There exists
a
of
capillaries.
heterogeneous
dividual wall substance
4.
of
1). Of these phenomena
(Fig.
distinguishes
The
is
why the cell’s
believe this
show
place
we
the
dia-permeation.
He
2.
is
(Incontestably,
Fig.
1.
carboxyl
higher degree of
a
permeability from
three
place,
by him:
tinguished
with
not
those cations that
Manegold (73)
In
water.
influence of NaCl and CaCl2
the facts.
permeable. If
soluble in water, permeate
when the film contains
true
a
while the Ca
soluble,
1.
low in the presence of cations
substances,
the
containing
down
properties
shall find Na within
we
brane. Now this Na membrane will show
be
the
insoluble. The other way round, the Na salts will be
perme-
water, whilst
protoplasm in
will
are
to
There
is
a
wall without
showing
heterogeneous
wall
a
different
with
capillaries,
each in-
degree of permeability.
capillaries.
8
The
of
permeability
the
first-mentioned
upon the relative volume of the
emanating from
In
the
must
the
enter
can
boundary between
a
pass
In
picture.
the
of wall,
the
mainly
depends
shape and the forces
capillary wall likewise playing
the
of the second type
case
wall
the
capillaries,
important part.
an
molecules
permeating
media, when surface phenomena
two
of the third
case
the relations between the components and their
of membrane,
type
nature
are
important.
The
mixed form of membrane shows all
three
previous groups. (Here, it should be stated that electric phenom-
ena
are
treated
not
regarded
The author’s
the
of
mathematical
that
a
the
shall
we
diffusion
by him. If
have
be
to
constants
shape,
of the
findings
permeating
In
are
few
a
will
physiological
of
no
simple
avail in
cases,
have
to
be
membrane.)
the
the
of
case
permeation
etc.
substance, but everything else indicates
membrane
physiological
established
that these varieties
of the
given membrane may be computed from the diffusion
of the
constants
so
forms
protoplasmic membrane.
constant
the
by him,
complex
less
as
the characteristics
we
are
belongs
to
ever
acquainted,
arrive
other
amongst
fourth
sound
in each of the wall substances,
of the
the
to
at a
type,
computation,
things,
with
the
the relative volume,
capillaries and the surface concentration of the
substance.
A
membrane
described
Fig.
15,
that
the
in
3).
There
These
watery
by
stable
affording
by
are
protoplasmic
formed
may be
1.
model
papers
an
films
two
membrane is
autocomplex
have
the
to
Jong
de
arguments
in
be
and
support
regarded
coacervate
power
to
of
Bonner
of
as
a
separate
a
3-
the
been
(14,
theory
film'which
phosphatide:
two
media.
Fig-
has
potentialities,
many
Bungenberg
Fig. 4.
miscible,
9
There
2.
between
analogies
are
of
amount
of
water
these
the action
of
electrolytes
and
coacervates
that
the
on
exercised
on
permeability.
is
Zone II
Zones
I
fatty acid
An
1.
2.
of the lecithin
molecules.
electrolyte
varied in
or
effective, electric
the
hydration,
In
two
of the
lipophile
Bungenberg
papers
phosphatide
If
coacervates
sensitizer is added
a
between
the
The
of
structure
different ways
substance may
water.
found between the
are
(vide
act
this
§ i).
on:
attraction,
the functional activities
to
three
organic
an
the
the attraction
3.
hydrophobe;
be
can
the ions and the
sensitizers
are
chains
membrane
hence it contains
hydrophile,
and III
of sensitizers
likened
are
will
become
growing tendency for the latter
them
to
soap
The
coacervates.
soap coacervate, the interval
smaller
and
there
line themselves up
to
31) refer
within the membrane.
by
amorphous
to an
molecules
groups.
Jong and Saubert (30,
de
will
be
a
parallel. This will
ultimately result in the formation of bi-refringing myeline tubes. The
more
the
the
will
be
is
coacervate
of
more
its
directed, the less
sensitizer is
a
for
permeability
the membrane described
action of the
CaCl
2,
of
be
tri-complex
brane
(Fig.
example
traced
opens
model
is
acid
and
phosphatidic
If
very dense.
between
i.e.
contain,
the lower
contend that,
the
from
new
up
in
the
de-hydrating function of
the model.
The
potentialities
for
discovery
the
mem-
4).
the
Now,
will
They further
water.
for
directly
systems
it
water
the membrane,
by Bungenberg de Jong and Bonner,
electrolytes,
cannot
found within
the
built
from
up
Ca
ions.
three
This
lecithin,
components:
film
will
be
presumably
replace the Ca ions by Na ions, then the interval
we
particles
and,
the
correspondingly,
permeability
will
increase.
By
the
means
of
comprehensive
erythrocyte
wall,
are
molecules
Bungenberg
This
model is
biology
(Fig.
expected
to
Jong and
de
play
a
an
the
of
phosphatidic acid
Saubert
exceedingly
nature
model in which
membrane.
important part in
5).
In the latter three models,
the membrane.
suggest
regarding
(ioo) develops
substituted for the chain of
protein molecules
of the
researches
Winkler
There is
referring
to
an
there
are
electric
as
two
well-defined sections of
well
as
an
a-polar part.
the forces that rule the condition of the
We
complex
as complex and symplex relations. We propose the expres“complex relations” for the forces in play between the ionized
systems
sion
groups.
“Symplex relations” is
a
term
coined
by Willstatter, by
10
Fig-
which
should like
name we
to
5-
indicate the London-Van
Waals
der
forces.
Antagonism of the ions.
§ 3.
It is
conceivable
of
complex
coacervation
distinct
3
useful
as
strongly
can
be
Co(CN)
salt
to
(inorganic
J
With
2
with
The
a
to
Thus,
complex
positive
nucleinate and
escence,
whilst with
)
1)
the formation
nH 0
2
coacervation)
->
Gelatine-arabinate. nH 0
2
>
can
biocolloids
coacervation,
will form
with agar it
be
no
interposed
2
will
show
in
a
series,
great
coacervate
slight
coacervation will
opaloccur.
which shows
thus: nucleinate
of the
complex relations,
amylum solubile > glycogen.
(en) 2 } 3](N03) 6.
show
complex
a
steadily increasing sensitiveness for
[Co{Co(OH)
as
the
Hexol-arabinate. nH2 0
gelatine
arabinate,
decreasing intensity
arabinate > agar
to a
regarded
causes
become ill-
to
Hexol-cobalticyanide.
-*
“amylum solubile”
negative biocolloids
points
solutions
(complex coacervation)
respect
variations.
solutions,
true
as
that
complex coacervation)
Ca(N0 3)2 (autocomplex
CaCl
(12).
sols
The fact
(13):
Gelatine-chloride + Ca-arabinate
+
be
hydrophile
ions
and colloid
Ca-arabinate + Hexol-nitrate
+
regard
detected in
true
+ Hexol-nitrate
6
+ KN03
to
hydrated
Complex coacervation is
“insoluble”
an
K
polyvalent,
boundary between
defined.
of
well
as
solutions
a
The
same
>
series
neutral salt, which
11
is
added
the coacervate,
to
decreasing
i.e.
the
From
quantitatively.
density of charge of the particles is
in the
seen
weight
from
increases
of
amount
colloids.
the left
of
density
hexol
reciprocal
to
hexol
have been found
equivalent
Na-nucleinate <
can
right.
to
Teunissen
weight.
by
form
to
the
as
cations
of
in this
vation with
manner,
amylum
<
26000
2264
coacervates
with
hexol number decreases.
will
sooner
a
given cation, increases
On
the
other
hand,
autocomplex
cause
given biocolloid than those cations that have
a
the
these
coacervation the density
charge of the
importance. The tendency of negative biocol-
higher valency
a
of
93)
of
autocomplex
reciprocal
92,
from
following reciprocal
Also, in autocomplex
loids
(34,
charge
at
The
Na-agar
1068
colloid ion is of great
equivalent
Teunissen:
Na-arabinate <
294
the
reverse
But also
decreasing.
say that the
number, arrived
may be likened
numbers
the
we
is
standpoint, the
charge of various biocolloids
needed
hexol-nitrate,
The
the
to
be
to
seen
light of electro-chemistry,
established the
gram of colloid
charge per
colloido-chemical
a
the
coacer-
lower
a
valency.
The
charge of almost
every
negative biocolloid
can
be reversed
by using different salts (92, 94). This reversal will take place
concentration
higher
cervation becomes
groups
belonging
plex relations
cations.
(arabinate,
sulphate,
ionized
between
those
concentrations
affinity
phosphate
colloids
colloid
and
for
the reversal
the less
groups
the
to
charge
by
sulphate
K
the ionic
among
colloids
> Na
affinity
amongst other
means
Li
> Na
things, when
alkali-cations.
represented
The
by:
respect
to
phosphate
increases,
colloids
thus;
> K.
standpoint, we can divide
complex colloid systems into uni-, di- and tri-complex systems
Departing
(Fig.
6).
this
is
of
> Li
with
the volume of the cations
from
define
of bio-
(lecithin, nucleinate), carboxyl colloids
conspicuous,
of
the other hand, the
as
to
charge
pectate)
becomes
reversal
with respect
decreases
com-
voluminous
been able
of the
a
of the ionized
polarizability
ions
at
coa-
and sulphate colloids (chondroitinagar). The influence of this polarizability of the
pectinate,
carragen,
defining
the
The
or
the colloid anions is very essential in the
(ionic spectra); he found typical differences
spectra of
On
flocculation
autocomplex
difficult.
more
to
as
By using sundry salts, Teunissen has
the different
colloids
point
electro-chemical
12
6.
Fig.
Whether
tri-complex
a
attractions
y and
x,
which in their
a
uni-
turn
z
is
be
to
be
may
The
di-complex.
or
tri-complex
system
it
Now,
is also
made
+
true
that
a
increases.
In
spite of the fact
hexol numbers
tendency
to
form
complex relations
w
depends
the
on
and
w
y,
the formation of
must
be filled
if
a
+ y.
tri-complex
reciprocal
the
>
z
degree of density
great
for
responsible
following condition
the
as
formed,
formed:
x
readily
be
can
compared with the attractions
as
(i.e.
to
be formed the
can
which the components
that
little
lecithins
density
marked
are
of
more
charged
are
by
high
they show
charge),
tri-complex systems. In di-complex systems
from the low
positive colloids are weak, as may be guessed
density of charge of the lecithin. This apparent contro-
versy may be
explained by the fact that the reciprocal hexol numbers
but
are
this
a
we
gauge
are
to
formation of
increase
its
ponents
are
Van
the
dhr
of the
of negative
excess
conclude that lecithin
than
tri-complex
purity.
egg
In
contrast
albumen,
Waals forces,
to
over
will
that of
with
gelatine,
a
more
readily
di-complex,
tri-complexes,
and
From
positive charges.
tend
the
like,
to
when
whose
the
the
we
com-
London-
existing between the long carbon chains of
molecules,
allows the
be
a
to
can
play an essential part in lecithin tri-complexes.This
coming into being of tri-complexes, whose existence would
incompatible from the standpoint of complex relations exclusively.
If the
protoplasmic membrane be indeed regarded
system of Ca, lecithin and
we
see
ions.
the action of salts
On
regarded
his
the
as
tri-complex
on
this membrane
as
an
must
antagonism of the
that the protoplasmic membrane is to be
auto-complex of lecithin, Bungenberg de Jong and
decided
antagonism of the ions,
of
a
assumption
an
collaborators
charge
as
protein, then, in the first instance,
lecithin
to
coacervates
their
concentrate
thereby starting
(16,
18,
from
26,
attention
the reversal
28).
These
on
the
of the
data
they
compared with the “potential milieu” of Artemia salina (L.) (2,8,66).
This showed them that
1
to
4
mol,
in the
region of concentrations of from
germination of Artemia eggs corresponds.
only
the limit of
13
to
certain extent, with the reversal of the
a
this
of
the
reason
working
lecithin) is probably
for
the
phenomenon.
(unknown
system
why the
Whether
1.
depends
quotient of the
exceed
place depend
(N0 )
2.
If
3
using
of adsorbed
amount
Nevertheless,
medium
to
adding
This
3.
From
this
we
4.
The
is
conclude
may
bearing
becoming
of the less
forecast
It is
of
these
out
directly
voluminous
probable
likened
However,
it
is
to
to
the
activity
an
We
anion of
anion of
§ 4.
The
it becomes
yet
3
.
phosphatide
we
in
K,A
this
by
that the
see
the
disperse
means
that the
grows
smaller
manifest
in
when
in low
even
dealing,
are
principle,
of the anions
valency
Under
known in
identical
decreases
on
it
is
more
biology,
conditions,
the
rapidly, than that
tri-complex system
a
may
on
be
considered
liken
the
activity
example alkali-salts
salts
on
to
be
or
phosphate,
cannot
here
charge
be
broadly.
organisms
on
the reversal of the
further expect
can
be
of colloids.
of the members of
alkaline earth
carboxyl
the influence of
a
greater than that of
or
salt
a
a
metals)
sulphate
possessing
salt
with
an
valency.
of
lipoid
to
a
to
of these
action
compounds,
between
and NaN0
ones.
acting
high valency
Overton’s
we
that the influence of salts
that
(for
may
low
a
detail,
possible
group of metals
colloids.
A,
complex cations,
facts.
the influence of salts
in minute
not
a
in
not
reason
and A.
that the
activity of voluminous cations
Though
charge,
importance.
exceptional position
also
2
adsorption
to
antagonism between Kj
of essential
K
salt
charge,
shall observe that
we
of
does
antagonism
Th(N0 3 )
of the
more
of the
antagonism
2
concentrations,
with the
require
reversal
a
the reversal
antagonism
charge of
a
are
change very little.
with respect
K,
K A.
will
Kj
we
cause
of
activity
as
understand
4
in the presence of
4
tri-complex
to
neutral salts
two
greater
than between
the reversal of the
K A
explanation
a
as
of the cation. For this
valency
a
autocomplex
an
(18):
at
s
salt
a
facts
Hence
10.
found
and NaNO
antagonize
we
the
on
2
as
For
water.
place when
concentrations
Jong
de
accepted
might be able
we
following
the number of
Bungenberg
2
take
the
must
U0
moment),
will
on
The
the first
be
to
germinate in distilled
not
antagonism
combined,
simple
an
treating the membrane
By
that
at
do
eggs
theory (the
too
of lecithin.
charge
membrane is
the
matter
organic
substances
theory puts
solubility
of
in
down the
on
permeability for organic
the membrane
importance
to
trace
coacervates.
lipoids.
Therefore,
the influence of
organic
14
substances
act
on
the
on
acids),
may also
we
of the
phosphatide
use
protoplasmic
The action
on
of
membrane
1.
the
non-electrolyte
the
lipophile
substances
oleate and
exert
either
phosphatide
non-electrolytes
as
fatty
model
a
depends
system
system,
coacervate
(17,
upon
upon
this
the stock model.
on
have been
33)
32,
shrinking
a
In
coacervates.
the
and
occurs,
film
Rosenthal
coacervates.
It
alcohol
exert
orientated
or
exercise
a
was
Fig.
electrolytes
different
the action
that
and
a
similar
to
dependence
speak of
a
from
that
Saubert and
of alcohols,
coacer-
and
propyl-
amyl-alcohol
(Fig. 7). Other
Fig.
It would be better
In
the action
number of soap
ethyl-
butyl-
coacervates
a
recall
we
water.
oleate
upon
urea
methyl-,
whereas
oleate
Jong,
research
7-
exhibit
consequently
be
may
for
investigating
actions,
on
opening action
respect
de
established
shrinking action
an
Bungenberg
(83) included in his
opening
or
this
permeability
ethers, ketones, urethanes and derivates of
atoms.
similar
a
upon
(amorphous
biological films cholesterin often
vates.
etc.
itself,
the action of these substances
Booij
chiefly
charge, presence of sensitizers).
Organic
exercised
compounds
the esterified
(i.e.
(32).
component of the
the condition of the
system
system,
of
these
facts:
numerous
upon
As
oleate-, stearate-coacervates,
non-electrolyte
a
2.
3.
coacervates.
lipophile part of phosphatides
non-
8.
upon the number of carbon
dependence
upon the
length
15
of the carbon
chain
that chain. Thus,
alcohol
slightly less
agent, while
tert.
and
the
to
that
has
butyl-alcohol
The influence of alcohols
likened
upon
the oleate
on
well
that
as
action
same
on
colloid
exercised
remember that the
must
electric
Action
from
uncharged
phosphatide
on
coacervates, sensitizers
of
the
that in
be
must
by
a
zero
renewed
opening action.
charge of the
The
occurrence
A
considerations.
cervation of
needed
At the
to
zero
find
we
phosphatide
practically
coacervates,
and the
the conversion of
The
employed.
choline groups.
at a
This minimum
is
distance
some
phosphatides
for
curves
un-
into
methyl-,
initial
opening,
ultimately followed
to an
is
we
important
an
There
coacervate.
the
with the
corresponds
coacervate.
of this minimum is explained by the following
given quantity of CaCl2 is essential to the coa-
phosphatides.
reverse
may be
(85), in his
the protoplasmic membrane
appreciable decrease of volume which
an
in
shrinking
coacervates
ethyl- and propyl-alcohol (Fig. 8), after pointing
show
weak
a
Saubert
(17).
Although
and
group
action.
inappreciable, when working
state
in the fact
pleasantness
as
phosphatide molecule contains
these groups is
the
oleate
acts
opening
namely the phosphate
constituent,
on
models.
on
an
hydrophile
shrinking agent, isobutyl-
a
coacervates
thesis, discusses the action of alcohols
as
as
acts
butyl-alcohol
sec.
so,
of the
position
butyl-alcohol
n.
the
On
charge
adding alcohol, the
of the
amount
phosphatide,
of
be
can
reduced.
charge, the effective attraction, exerted by the particles
in the coacervate, will be found greatest, hence the volume decreases.
Once the
zero
weaken and
point of charge is passed, the effective attraction will
the bulk
of the
coacervate
will increase.
Saubert measured the action of alcohols
phatide
coacervates
at
different
points
the volume of
on
of
CaCl
phos-
concentration.
2
this
theory, he found alternately an increasing and a
decreasing influence of methyl- and ethyl-alcohol, according to the
Following
is
degree of CaCl2 concentration. Propyl-alcohol
agent,
butyl- and amyl-alcohol
an
alcohol
on
the carbon chains and that
on
a
condensed system
When the system
action
also
grow
in
on
as
opening
an
depends
on
two
facts: the action
the electric condition of the system.
the
point of charge,
zero
condition will be
importance
always
shrinking agents. The action of
approximates the
the electric
on
are
density
of
then the
this action will
appreciable,
charge
of
the
colloid
increases.
As the
protoplasmic membrane is regarded
the
action
of alcohols
as
the
an
orientated
phatide
system,
often be
compared with the action of these compounds
colloid
systems.
Whereas
on
permeability
butyl- and amyl-alcohol
on
act
as
phosmust
orientated
shrinking
16
agents
amorphous oleate coacervates,
on
opening agents
as
is
on
centration,
find the former
we
liquid oleate crystals.
follow the Traubean rule
to
seen
alcohols
upon
erythrocytes).
to
(vide the haemolytic
action of
obtain
the
of NaCI
degree
liquid crystals,
increased resistance of the system
correspondingly
act
influence
An increase of the
which is essential
to
The latter
con-
causes
a
the influence
to
of alcohols.
We
shall have
fatty acids
that
reckon with another feature of
to
of the lecithin.
Rosenthal’s
methyl-, ethyl- and propyl-alcohols
act
influences.
Now, if we take K-oxystearate,
shrinking action caused by propyl-alcohol.
a
hydrophile
considerable
ricinolate
(oxyoleate)
the
ethyl-alcohol
identical
group
into
action
the
will
be
the
will
shall be faced with
observed if
a
carbon chain is of
we
of
curve
practically
shorten
An
constant.
the chain
of
Kand
adding methyl-
on
a
the introduction
propyl-alcohol
remain
upon
causing shrinking
Hence,
decline, whereas
volume
will
we
lipophile part of
Thus,
importance.
are
the
(83) confirm
opening agents
as
K-stearate, whereas butyl- and amyl-alcohols
of
importance:
experiments
fatty
Propyl-alcohol will show a shrinking action on K-laurate,
ethyl-alcohol shows an inappreciable shrinking, followed by opening,
acids.
whereas
It is
we
must
in the
of
our
when coming
dealings with the action of alcohols
ascribe this
protoplasmic
direct action
a
this influence
the action
to
§
When
an
is felt, lie close
on
the
Traubean
biological objects,
the
fatty acid chains
of course, the
eventuality
If the concentrations,
at
which
together, then this may be attributed
upon the electric
substances,
there
explanation of the facts.
important,
no
The elective
2.
A
3.
Adsorption
4.
(excepting,
metabolism).
on
on
condition of the membrane.
recording the elective permeability of the plasmic membrane
certain
most
1.
the influence exerted
to
membrane
across
Permeability.
5.
for
open up.
believed that,
generally
rule in
will
methyl-alcohol
sieve-
or
Electrostatic
the
four
doubts others
solubility
filterlike
the
at
are
mechanisms
Though these, in
within
may be
the
cell
our
likely
opinion,
to
offer
are
the
forwarded.
membrane.
action.
protoplasmic
attraction
or
surface.
repulsion
between the substance
and
membrane.
Each of these four theories has found staunch supporters. Overton
found
a
correlation
in oils, and their
rate
between the
of
solubility of organic compounds
penetration. His lipoid theory suggests that
17
they
be
can
dissolved in the
lipoid components of the protoplasmic
membrane.
Ruhland
notes
great influence of the size of the molecule upon
a
permeation of the compounds
the
molecules have
(ultrafilter
slower
a
Traube observed
that the
plays
membrane
as
this is
the other
on
obliged
look for
to
by
combination
the first
theories
surface.
by
negative
a
by it.
facts,
numerous
comprising
as
that
so
many
facts
place,
in
lipoids
activity, according
Meyer
cules
be
well
as
at
first
has
the
as
adsorption-affinity
adsorbed
his
greater
onto
Waals
stances
Collander
ceratophylla
an
rate
a
the
for
many
An
at
of the ultrafilter
concur
to
organic compounds.
a
quicker
in
substances
are
the
London-
wall
sub-
rate
factor
certain
are
for
their
than
lipoids and the
one
would expect,
even
the
the
sense, but
a
molecules whose size
rate
of
permeation.
lipoidfilter theory,
is
All of
which is
not
recognition of the fact that both
The
been examined in many
45, 5L 59, 60, 61, 62,
Generally, there
in
lipoids. On the other hand,
important in permeation.
has
solubility
theory (who prefer Beggiatoa for their
establish
theory in the proper
principles
lipoid
exception is constituted by the small-sized
that permeate
only deciding
these facts
Here, the
permeating and
correlation between the
experiments) admit that there
not
Hence,
HaNsteen Cranner
permeation.
judging from their solubility
supporters
mole-
the
a
fight in favour of the adsorption
partisan of the lipoid theory (56). Winkler’s
to
permeation.
molecules
chains.
(37, 38, 39) has measured the permeability of Chara
Wallr.
obvious
of
The
capillary
homologous
colloids.
sympathy.
forces between the
decisive
are
and the
with small-sized
model reconciles these theories with each other.
der
lipoid
the
putting up
theory (55), becomes
Van
and
adsorption
Traube’s rule, increase in
to
intensely
of narcosis
after
also,
that the
(76) maintains that many narcotics
cannot
theory
remark
we
only be divorced from each other with difficulty.
can
solubility
a
those
by the charge of
attracted
are
possessing
by the fact that the
repelled
are
supported
a
out
determined
hand,
permeating
possible.
In
is
ones
than
rate
protoplasmic
is
Anions
(Michaelis).
cations,
of the
quicker
a
borne
the
on
electrolytes
Each of these theories is
is
Large-sized
than the smaller
Compounds
part.
at
permeate
and
of
permeability
membrane,
one
living cell.
adsorption-affinity
strongly adsorbed
are
The
the
permeation
important
an
affinity
small
a
former
the
also
adsorption-affinity
having
into
theory).
substances
great
of
rate
permeability of organic
organisms
(7,
37,
38, 39,
63, 64, 67, 81, 82, 84, 86, 87, 88, 95,96,99,101).
2
18
In all
of these
researches
we
both
are
confronted
with
the
theory (or adsorption theory) and the ultrafilter theory. The
of the membrane, shown
for the
bear greater witness
in
is
theories,
the
lipophile
brane.
well
as
The ratio
either the
lipoid
in
countered
of the
and others
As
cells
from
simple
to
of
The
electrolytes.
make
—
establish
the other three.
our
to
problem
a
difficulties
others
anions,
to
en-
cations
to
very intricate
one.
understand the influence of salts
between tins
relationship
a
Moreover, it
Efforts
with
one
More
is
plasm
reconcile
to
the
another
not
are
under
principles
theory (Nathanson)
mosaic
favoured
as
a
of salts
consideration
and the emulsion
theory (Clowes)
solution of the
the action of these substances
biological point of
with
view. Most
the
on
that
compounds
conspicuous
is
decreases under the influence of CaCl
of the
permeability
are
the influence of salts
Stewart and
.
(88) have observed that the permeability
is greater in
of
is
noteworthy that these salts do
ethylene
Gellhorn
glycol.
of
buffered
to
staining is accelerated by
The
MgCl .
or
2
considerably
counteract
Organic
we
are
less
MgCl
Finally,
advance
of
CaCl
2
to
not
water
or
influence the
(50) has submitted the
solutions
of
neutral red.
treatment
makes
2
of it is needed
oppose
salt,
finally
The
rate
of
with univalent salts
coloration
impossible;
NaCl and KC1 than
to
.
substances,
then
an
addition
presented with
lated and
for
2
It
Strongylocentrotus
placing them in a solution of nile-blue
eggs
of Arbacia eggs
while
Jacobs
solutions of NaCl and KC1 than in solutions of CaCl
seawater.
permeation
from
interesting
2
and
—
problem.
permeability. Both NaCl and KC1 stimulate permeability,
this
the
—
important than the permeation of substances into the proto-
membrane, when dealing
on
and
principle
that the
appears
permeation
strongly influenced by cell metabolism (71, 89, 90, 91).
is
of
prevalence
given object.
a
permeability than the permeation of the salts proper. It is far
on
a
principle in
being permeable
both
to
between both
permeate
determines the
two
rule, it will be found easier
a
can
by the charge of the membrane, is especially
permeation
some
again
former
the ultrafilter
or
the
—
difference
great
through
through the hydrophile parts of the mem-
as
The action, exhibited
important
organisms present another picture.
model the
longer apparent. Compounds
no
plants and animals
lipoid than of the ultrafilter principle,
other
some
Winkler’s
Most
principle.
one
of the
though Beggiatoa and
Again,
by the object under observation, is deciding
of either
prevalence
lipoid
structure
too,
a
influence
difficulty:
again it will
in certain
be
instances
a
(4, 5). But here
permeability will be stimu-
permeability
now
checked.
substance may
move
in
spite
of
a
19
difference in concentration.
This
part.
is
permeation
meation under the influence of
a
model
be
by Teorell (91)
when
passive;
the cell
of
plants
virtually
depends
acid) is
line
a
the
can
the Donnan
the
through
Hence,
too,
continually
substance present
establish
to
salts
respiration.
draw the
to
of
taking-in
cell
permeation,
an
do
we
between adenoid
permeability (Hober)
permeation of a given substance
hairs
root
think
not
it
permeability
and
physiological
or
permeation,
the
on
possible
(Overton)
(e.g.
be substituted for
can
of per-
difference in concentration. From
a
learn that this
we
active
an
adenoid action
an
(for instance ions tending
Thus,
equilibrium).
metabolism takes
as
to
given compound
a
formed within the cell, it
outside
cell
Here,
referred
“passive”
may be
passive,
though, in reality, cell metabolism will be chiefly responsible for it.
And
He
as
thus
found
enter
we
the
realm of
Straub’s
difference between the
a
sort
of
this
difference.
equilibrium,
secondary
however,
two
as
a
is
very
while
occurs
an
acid
without
further
of the energy:
1.
of the membrane,
no
(root
2.
than
a
cases,
There
are
into the
Respiration
ions will diffuse
whereas within
egg)
special
some
diffuses
of it.
out
a
maintain
to
hairs).
Oxygen
case
hen’s
more
in
interest;
2
boundary. In this
be
a
not
required
difference will
(mostly C0 ) will pass
in the surface
the pores
this
important functionally
potential suppliers
cell,
of energy is
source
Generally,
phenomenon
it
This, he maintains, is
respiration.
90).
concentration of the
yolk and that of the white of an egg (experimenting with
and put this down to
(89,
researches
osmotic
through
the membrane
trans-
portation of electrons will take place.
In
both
if
occur
membrane.
rate, while
is
he
cases
centration
Into
a
large
a
(harmony),
out
as
is
at
amount
added,
ions
are
plays
This
will
much of the
concentration
the vessel.
many
Hence he
to
sulphuric
Many
it will be
occur
will
varies
situation
K
is
much
ions
than
of the
surroundings
sea-weeds).
establish
concludes:
considerably
understood that in the
between the
If
itself.
electrolytic
one
“kills”
for
the
As
different
living cell great
composition
(vide the strong
a
slow
acid diffuses
cations
more
a
concentration
stable
a
of the
at
solutions. The
the membrane,
vacuole sap and the
equilibrium
a constant
gives rise
con-
regularly through
equivalent
mobility
important (though incomprehensible) part in this.
(Michaelis),
found in
the
differences in
sulphuric acid is poured
taken up from
an
differences
as
than outside
the former, within
ions
ion diffuses
an
of salt solution of
point
while
or
vessel
the vessel.
which
stronger inside
Li ions
substance
porous
present outside
models in which
develops
a
K
of the
concentration
protoplasm,
true
20
If
is
energy
condition
continually
(harmony),
produced
at
from
diverging
this energy is made up of electric circuits,
in.
set
At any
These
of
theory it
is
as
Can the
plex
the
on
depend
composition of the medium. From
protoplasmic membrane be regarded
membrane
should
stimulated others
(14,
be
15)
regarded
further
to
the effect
to
as
an
biological
that
the
will
cause
point of charge
of the CaCL
water.
If the
for
this
and
cells,
found
for
a
Ca(N0 )2
3
a
This is
this
a
influence
Ca-autocomplex
NaN0
opened
up, by
importance of making
salts in
The
to
numerous
3
.
a
and
by such
zero
in-
phosphatide
a
definite
with
theory.
to
a
Moreover,
exist.
strong
on
structure
NaN0
3
as
does
original
these
be
work
reached
De
Haan
membrane.
hence
antagonized,
confirm
a
the
to
not
concentration is
of the
experiments
of
was
the membrane.
which is
comprehensive
while
membrane,
concentrations
Strong
Weak
have
3
protoplasmic
rule
of Allium
water
the
Co(NH ) 8 C13 (luteoCl 3)
the
very
great
study of the behaviour of
concentrations.
minima
produced in De Haan’s permeability curves are
correspond with the concentrations at the reversal of charge
of the
phosphatide
system
we
are
first
the action of the salts,
on
the
and
the
further increase of
a
of neutral salts,
concur
of lecithin
His
at
subsequent
a
permeability for
to
exerted
the
to
phosphatide
(53) defined the influence of salts
the
until
not
be
a
found.
permeability.
valency
action
opening
ascribes
said
of
to
be ruled
decreases.
water
increase
he found
the
on
weak concentration;
an
on
the facts
influence
an
cause
of
be
De Haan
3-valent cations
permeability
at
will lead
water
but
attained,
has
direction.
of water;
amount
certain concentrations
assumption,
2 or
expected,
at
permeability will
condensing
will
concentration
permeability for
concentrations
salts
be
2
membrane, then,
cepa
com-
system,
research in this
decrease of the
a
minimum will
a
crease
with 1,
a
protoplasmic
autocomplex
An increase of strength of the CaCl2 concentration in
coacervate
minima
as
saying that the theory forwarded by Bungenberg
Bonner
Jong and
On
occur.
the membrane
on
system?
It goes without
de
expected,
directly inferred that the membrane consists
be
may
be
to
If
occur.
then electro-osmosis may
phases.
two
§ 6.
stationary
a
may
differences in the ionic concentration will
rate
differences,
(mobility of the ions) and
his
boundary,
a
equilibrium,
to trace
ruling
charge of the protoplasm.
Wakkie
permeability.
is
obvious
employed by
Bungenberg
(23), when plasmolyzing
It
de
Jong,
De
that
Haan,
De Haan
Spirogyra cells, measured the
21
velocity
of the
trations
of the salt.
attain
to
the
protoplasm in the electric
Using
which
CaClj
by
value
of
means
be
may
electro-endosmosis.
they
the
on
the
assume
As
adsorbed.
haviour,
This
raised under the influence
potentially
found
they
the
(vide
Under
at
exhibits
the
on
to
no
vesicles
U0
sequence:
2
With
2
reappear;
the
be
can
rule
valency
action of salts
on
used
they
means
a
is
be-
electrophoretic
behaviour of the
(57), is due
droplet.
layer, adsorbed
employed by
are
may
be formed
Ehrbg. This phenomlesion. Two factors
to
are
conspicuous proof
regions of concentration
Neutral salts show
Ca <
Na.
the
hexol-nitrate. Salts
This
of
counteracted.
said that this
charge of
exert a
a
vesicles
salts
can
antagonism
§ 1). The
(vide
same
rule
in
on
the
butyl-
make
them
gives rise
applies
membrane, reversed by
similar action
the
of
in the membrane.
hyaline
Other
follow this
action
powerful
phosphatides
formation
hence it may be
continuous
sur-
negative
conclusion.
the existence of
to
solution
a
of arabinate
appear. Concentrations of the cations
hexol-nitrate,
alcohol
the
on
be
to
the influence of alcohols
< luteo <
points
the
to
charge and the elasticity of the membrane. His
of the Traubean rule.
ion
this
film
the existence of a
Hartkamp
by
it: the
findings regarding
U0
at
a
surface of Paramaecium caudatum
essential
where
action
an
condition,
which
morphological
assume
arrive
to
investigated
enon,
answer
permeability and
vastly different conditions hyaline vesicles
the cell
are
§ 2)
of this
onto
though entirely different
protoplasm,
them
on
hydrophile colloid with
a
droplet
Danielli and Davson also
onto
by
representative
film
not
re-
charge of the protoplasmic membrane. Indeed,
coacervate
though
unable
the
at
phosphatide system of this membrane
model
a
phosphatide
concen-
were
extrapolation, they found 500 m.eq.
Likewise,
rounded, through adsorption,
charge.
they
Arriving
.
present controversy between the action of salts
the influence
varying
at
theirs,
point of charge with CaCI2
zero
versal concentration
of
field
apparatus like
a
the
means
of a
charge
to
to
of
complex
system.
Saubert
plasmic
(85) investigates the influence of alcohols
membrane and
colloid
models.
different lines of conduct in their action
(vide § 4). Firstly, they
may
chains;
may
complex
support
secondly, they
each other.
influence of alcohols
permeability
have
some
on
semblance,
may
phosphatide
influence the electric
Saubert
of Chara
on
on
the protofollow
two
coacervates
change the interval between the carbon
These
components.
Alcohols
two
powers
finds
some
phosphatide
ceratophylla.
though
n.
may
condition
either
semblance
coacervates
between
and
that
Generally speaking, the
propyl-alcohol
acts
of the
oppose
on
on
or
the
the
curves
Chara cells
22
as
shrinking
a
the fact
by
agent.
centrations than
This
is
as
an
ascribed
The
charge.
plained
act
on
phosphatide
on
divergence
of the
The second
that alcohols
point of divergence
the
cell
living
to
difference in
a
shrinking influence, he continues,
action
the
on
direct
measurings
(ioo)
Winkler
manner.
of the red blood
or
the
con-
to).
:
density
be
must
oppose
ex-
themselves
behave in
charge, erythrocytes
puts the
different
a
whether the membrane
question
should be
corpuscles
with
experimenting
of
of the
i
charge of the complex system.
Whereas the cells under review either exclude
to
furnished
(the ratio being
coacervates
by him
is
much lower
at
regarded as a complex. By
erythrocytes in isotonic and hypotonic solutions
discovered a maximum resistance existing at the isopoint. The charge of the erythrocyte is influenced by neutral
sugar, he
electric
salts, according
neutral
on
three arguments
sistance
continuous
to a
salts, according
is
to
valency rule. Resistance also depends
in favour
are
determined
by
of
and that,
erythrocytes,
of
From
attraction.
These
valency.
theory according
a
electric
facts Winkler concludes that the
of the
rules
parallel
two
which
to
the
re-
following
represent the membranes
stromata
they determine their
simultaneously,
stability:
the isoelectric
the ionic
the
same
points
in
(vide
spectra
stromata
and
the membrane of the
Hence,
and
complex
subsist.
an
cannot
concur;
erythrocytes
in various
of
salt
is
of cations
valency
2
up
increased
the
Likewise,
Ultimately,
regarded
both
as
a
to
In
salts
(hypertonic
surmises
pn
an
according
were
found
to
From
of
a
correct:
a
show
show
ranging
such
negative
valency
following
this, it is
negative
the
a
hypotonic
the
to
destroyed
haemolysis).
con-
colloid.
stromata
are
tri-complex system of lecithin, stromatine and
corpuscles
may be
lined
A
up
parallel,
great many
with
properties
their
of the
explained when closely inspecting this
model of the membrane. Both the
An
resistance
autocomplex
cation.
The lipoid
molecules are
paraffin chains towards the exterior.
red blood
Its
11.
membrane is
a
this
complex,
a
the presence of pn 7
by cations,
erythrocyte
cluded that the membrane is
be
is
erythrocyte
electrolyte medium (NaCl) the erythrocyte shows
solutions
rule.
out
and
erythrocytes
stromatine.
charge, ranging from pn
to
erythrocytes
semblance.
between lecithin
a
and
stromata
sequence;
the stabilities of
great
In
stromata
§ 3) of
pores
and the
lipoid theories bear
system.
explanation
can
now
be offered for the changes of
permeability
23
under the influence of
electrolytes,
hypertonic
and
By
in
Bungenberg
tialities
of
(Fig. 5).
simple
The
the
in
so
row
be
can
of
lipoid molecules
membrane,
the potenincreased
considerably
potential membranes is very great for the
many
the
of the constituent factors may be varied:
and the ratio of the
nature
permeability for
amount
different
of
is
water
lipoids
essential.
are
defined, amongst other things, by
cholesterin present
which,
naturally,
may
vary
cells.
The ratio of the cations
4.
haemolysis.
a
protein may vary.
Both
3.
membrane model
that
reason
antagonism of the ions, the
Jong and Bonner
de
The number of
The
2.
this
the
the reversible
protein molecules for
substituting
the
1.
haemolysis
greatly affects
the
stability of the
mem-
brane.
Winkler’s
membrane model shows
It reconciles with
1.
The
2.
differences
reduced
in
the
to
another the
one
of
permeability
differences
important properties:
two
existing permeability theories.
organisms
exhibited
or
cells
can
by the tri-complex
be
com-
ponents.
When
with
comparing
under the
developed
great variations in their
The lecithin
another the three membrane models
one
direction of
Bungenberg
de
Jong,
we
note
properties.
uni-complex (Bonner) is chiefly governed by
sym-
plex relations. Of the three models under consideration this particular
one
salts.
at
is
most
greater the
(The
the reversal
the
of
When
the
of
relations
exceedingly
.)
The
tri-complex
to
contain
cations
becoming
are
In this
case,
i.e.p.
will be
found in
(Saubert),
systems
to
that
grow less.
are
systems
the number of
At this
exchangeable.
The
important.
more
important in tri-complex
active part.
very
2
7-
=
sensitivity for pu and salts
membrane will
greatly influenced by
of lecithin, the lower the concentration
with CaCl
pu
It is
changes.
pu
purity
proceeding
next
complex
to
charge,
neighbourhood
shall find the
an
sensitive
where
we
point
Here,
latter
are
protein plays
potential membranes is
great.
It goes
many
without
that
saying
potentialities,
many
we
are
to
regard these models
intermediate
stages
being
as
also
so
con-
ceivable.
§
7.
Introduction
the
to
experimental
part
of
this
research.
This
research aims
plasmic membrane. If
at
we
investigating
want
to
the
structure
of
the proto-
ascertain whether the membrane
24
is
complex system,
a
we
may
let ourselves be
guided by the following
criteria:
The neutralization of
1.
cordance
with
rules
The influence of neutral salts
2.
At the
3.
we
point of charge,
zero
are
find
to
Disintegration
4.
As
the
some
components of the
On the
the
arrive
is
the
at
the cell.
we are to
Hence,
unexpected
an
of the
as
manner
salts
due
being
pound
is
it
of
as
to
a
no
an
plexes
will
be
a
in
a
to
establish
any
on
cell.
at
all.
lines,
being
an
(i.e.
shall
we
cell
We
manner.
of the
we
do
the
regarded
not
we
We
a
process
in
a
surroundings.
biological
which
the
It is
the cell
changed
possible that,
an
permein
interpretation
nature,
are
be
at
yet
of
they
our
will
disadvantage if
know what
shall be able
of
substances
to
catalysis,
com-
observe
of
com-
be
very
we
want
phy-
permeate,
if
the influence
even
leaves
We
findings
membrane along
protoplasmic
stead
the model
explained from
a
a
course
the presence
assume
be
can
internal system.
several times in the
can
surface
shall
we
The membrane
membrane) in respiration,
as
we
cannot
and medium
which
moment
upon
the
hypothetical
nature
as
action
Although
Nevertheless
this surface
latter,
to
observations
our
straight-forward
siological
data that
though
cell in different
great
presumption
somewhat
the
97).
by closely examining
substances,
and likewise
valuable if all of
or
the
cell wall.
change of the membrane, the influence of
within the
of
(23,
where
active part.
and
faced with this
research,
re
study of the membrane model
elect
we
play
to
may be ascribed
shall be
our
a
to
solid
membrane.
their results
by
organic
ability will be noticeable in our object.
of
a
of indirect
being the region
to resort
long
boundary is expected
influenced
chitin
knowledge
within
protoplasmic
to measure
principle
study
none
measurings of the charge on
charge is a reliable gauge when
the
even
explain the behaviour of the
we
process
further
themselves of the membrane
position
in
without cellulose,
some
only by availing ourselves
shall have
we
cells
molecules,
studied direct from
surface substances may be adsorbed that tend
The membrane
contact,
few
a
may be
protoplasm that is enclosed
changes
a
con-
water.
of but
charge,
derive
may
not
structure
the
measure
often be in
a
membrane from
charge in
it
Hence,
we
But
protoplasmic
alter
to
a
the
investigating
other solid wall,
dealing with
exhibit
must
field.
thickness
a
excepting
When
the cell’s surface.
charge
of
amount
the electric
the membrane has
living cell.
the
on
ac-
valency.
when effective attraction is strongest,
minimum
a
in
of these criteria,
of
rule.
valency
tinuous
if
complex relations by neutral salts in
parallel
two
though the
no
room
for
25
Also, in respiration
permeability.
in
one
way
This is made
1.
the
reducing
3.
the
as
into
(89,
(sugar and similar substances) permeates towards
the first
resp.
systems,
membrane
of the
itself
is
chain
in the
aggregated
are
and
oxidising
an
two
On
hand,
the
other
lines
physiological
be
may
we
referred
when
to
Research
is
select
and
this
It is
object
our
organic
defining
to
factors
which
mutual
as
far
are
uniin
salts
Organic
factors.
essential
are
or
a
compounds
we
are
may
has been established.
a
coming
an
hydro-
one
chemical)
formed
to
saying
this.
of the constituent
we
shall
influence exerted upon the third
instances the first
individual
are
As
no
and second groups, too, will
between the
particles
on
object these factors ruling the situation,
complete
available,
erythrocyte
This is
salts
must
groups
theTraubean rule,
we
data
(either
furnish
cannot
biological
a
precise
membrane. An endeavour in this direction
any
The
into
Thus,
complex.
It goes without
The influence upon the interval
different.
is
the
subdivided
tri-complex
on
influence either
concerned with
in
attraction,
the
of sensitizer present.
In each
1.
be
ionized
be
of factors
groups
forces
can
the various
across
of
the influence of the
Waals
will
of certain
The results
hydrophobe part of the membrane depends mainly
amount
be
this
groups
that there
when
non-
cell metabolism.
influence
three main
di-complex (cf. Fig. 6, § 3).
group; in all other
the
along
“physiological”
processes.
particular have much influence
be involved.
in
on
the condition of the
to
between
proportion
Generally,
find that
as
der
Each of these main
attractions
such
not a
that
a
for the condition of the membrane: electric
groups.
and
life
that there
hydration and the London-Van
in
the
trace
its
on
the model,
on
catalysis.
permeability
shall then compare with
we
well-known fact
phobe
exist
system
this:
in order
compounds
substances
a
responsible
the
reducing
often confronted with
are
conducted like
investigation
same
and
oxidising
surface
as
permeability of the membrane largely dependent
We
of
membrane,
90).
Only the latter
salts
involved that,
are
with the substrate.
contact
interior,
the ferments,
as
ferments
many
come
must
possible:
the substrate
as
2.
another,
or
be
borne
regarded
(also
specific
out
in
region
wall
is
a
might
powerfully
description
of
be undertaken:
sensitized
system.
by the haemolytic action of all alcohols which
as
NaCl)
of
(100)
may
colloido-
or
a
neutralization of the
the
membrane will
concentration).
symplex relations.
subsist
Furthermore,
(at
the
least
in
In
a
wall will be
26
fairly firm;
The
2.
stromata
Amoeba wall
NaCl and
remains
is
that exerted
It
distilled
in the
the
that,
appears
(24, 57)
unknown
on
ranks
destructive
with
contrast
This
of the
case
sensitized.
membrane
is
between Ca
haemoglobin.
while the
only
and
when
possible
the
colloids)
are
erythrocyte. The influence
it
though, presumably,
far
as
membrane will
action
from
much less
will
differ
from
erythrocyte.
close
action
presumably
water.
(existing
powerful than
of alcohols
3.
in
relations
free
prepared
is
destructive
a
exert
intact
complex
more
KC1
be
can
(35, 36)
of
the wall
as
the
to
all
Here,
(also
too,
of
in
placed
when
Salts
Paramaecium
concerned,
(formation
alcohols
injured
be
Amoeba).
is
erythrocyte.
distilled
NaCl)
exert
the
note
we
The
vesicles).
(in
water
stabilizing
a
influence.
In the next chapters we shall be concerned with two objects,
namely the pollen of Lathyrus odoratus L. var. Pinky and bakers’
yeast.
we
It shall be
shall be
be
to
on
the
various
the influence of
some
the action of alcohols
be
faced
with
to
the
tackle the
as
us
(the protoplasmic
complex system) when proffering
compounds
to
carrier.
For
the
influence
of
when
ascribing
not
Chapter 111, § 4),
(cf.
of the
present
Even
complex
between active
or
must
we
relations,
restrict
an
complex and symplex relations,
important part
in
so
as
it is
many processes
CHAPTER
EXPERIMENTS
§
1.
we
may
such
group
ourselves
successful examination of the membrane proper from the
of
many
changes of the membrane like
fermentation
those between enzyme and substrate
colloidal
the
ruling
of life.
processes
neutralization
a
a
differences
on
problem from various angles;
theory accepted by
regarded
of
explanation
substances
task
guided by
membrane is
an
our
as
and
to
a
standpoint
the membrane that
plays
of life.
II.
WITH LATHYRUS POLLEN.
Introductory.
In
some
1934 Bungenberg
experiments
germination
program
of
de
regarding
Jong
and
Lathyrus pollen.
along which further research
that the material
Henneman
the influence
In
this
was
of
paper
to
(25)
published
neutral salts
be
they
on
the
developed
directed “in
a
order
might be compared with the behaviour of colloid
27
models
(mainly
studied
by
This is
cm
quantity of pollen is placed in
3
salt solution.
Ba > La
centage of
Na
>
This
when
rights
plotted
transition
from
be the
same
to
If the
3.
for
ion
of
shows
germination.
these
plates
solution
25
§
will
the
con-
advantages:
graphically
They have equal
the abcissae,
to
salts.
belong
to
the
by but
family, the transition
same
parameter, which is log.
one
point of each of the curves.”
remarkable behaviour:
(50 m.eq.)
liberate
readily
do
we
Ca
detoxify
to
ions.
the
Even
NaCl,
a
in
even
observe
not
warning against
are
the
happens
the
highest
depression of
a
of
use
agar-plates
2.5 m.eq.
a
CaCl
KC1 solution
resp.
2
of
m.eq.
2.
is
the
Measuring
The method
modified.
slightly
germination.
Bungenberg
employed by
3
Firstly,
we
of the salt
be measured. We shall have
to
strong solution of sugar
Off the flowers
water.
for each
to
and the
stamens
we
point
be
cm
order
3
In
).
keep
to
poured
this
out
off the
a
a
A
little
cut
we
operate
using
a
obtain
a
20
the
are
will
per
in
and
(taking
sugar
stamens
3
a
solution
which
—
is
tubes.
operated by hand, after which
droplet
liquid.
a 20
per
of this
The
cent
pollen
pollen
sugar
grains
solution
suspension
is
submitted
are
(
now
.05
(abt.
—
in
carefully
centrifuge
of
the
tube. Next,
centrifuge
are
flowers
containing
suspension
The latter
moderately
a
burst in distilled
carinae
cent
solution
cm
3
collected in
pollen
floral leaves
a
tubes
test
3
and
floral branches
we
These carinae
the
supernatant
again in
point).
back
into
brisk action of
of various
pollen
way
to
pollen grains
measured)
pollen.
shake loose the
the
as
Jong and Henneman
number of
a
prepare
cm
sucrose
de
(40 per cent)
solution
containing 3
20
per-
logarithm of salt
following
comparable
a
a
able
toxicity:
the
logarithmic x-distance, representing
various
They
be
germinated
plotting
plot arithmetically.
we
be characterised
concentration used
the
the
sucrose
logarithmically.
curves
(concentration)
The Ca
They
3
of
sequence
recommend
the level of the blank
S-shaped
range may
cm
the five salts used in their
to
following
has
It may appear that the
2.
been
active in low concentration are,
are
disadvantage when
at a
as
the
Ca.
>
procedure
The salts which
mixture of 3
a
germination obtained against
centration.
“1.
found
they
K
>
have
After 4 hours the percentage of
pollen grains is measured. With regard
experiments,
which
phosphatides)
years.”
by them;
applied
the method
A small
and 3
of
coacervates
in the past few
us
to
we
the
pour
suspended
cm
3
for each
dropped into each
28
of
the
tubes
test
which
hours
germinate.
to
the
pollen grains
testing
HC1
2
and
n
1
the
on
ultimately
of this is
cm
cane
passed
scope while,
established.
we
may
Though
that the
say
than
no more
of the
trying
When
we
pollen tube be visible,
The percentage of
grain has fully germinated.
in this
establish,
to
found that
the
salt concentration.
the influence of
manner,
arrived
curves
After
number
a
germinated grains is
germination is plotted against the logarithm of the
salts,
results.
Now
counted under the micro-
are
tip
a
fix
to
equal volumes of
dishes.
the percentage of
simultaneously,
have
We have been
the best
gave
into the Petri
pollen grains (generally about 250)
of
of
mixture
a
sugar
two
germinated
shall
hours.
two
then
for
the influence
we are to measure
we
We
left
percentage of the
of these
expiration
cent
40 per
3
hours
little time. If
a
fixatives;
many
.1
the
is
pollen
of concentrations of different salts,
series
a
shaken.
thoroughly
dishes where the
Measuring
grains demands quite
of
then
are
empty the tubes into Petri
would
at
some
reproduce.
not
Even the blanks
in
(exclusively 20 per cent sugar) showed variations
germination of from 50 to 90 per cent. At times KC1 chose to be
toxic in
a
reached
a
.003
it
of
strength
germinating
affect
solution,
n.
power
of the
germination; after
at
other
We
n.
.01
times
must,
rain the blank
was
low
(up
the
found
was
to
had
stabilize
to
The weather
pollen grains.
a
until the solution
not
therefore, try
50 per
to
cent.),
being high (abt. 90 per cent.) when the weather was fine.
By exposing the pollen grains for some time to a comparatively
stable
climate
would
(the
reproduce
of KC1,
CaCl
laboratory),
and
sucrose
like
considerably
vary
that
of
the flowers
Germination,
carried
instructions
discretion
a
are
when
be
at
constant
its
to
the
pollen
same
20
per
cent,
issued.
Nevertheless,
judging
the results.
select
our
no
appreciable
experiments
maximum. All
for
our
measurings
suspension
on
one
we
for
a
shall
have
if
example,
series
the
2
days.
are
now
solution unless other
of the alkali
and
is greatest
fully
experiments
sugar
For
influence
compare with another the
advisable
2,
shows
being kept in the laboratory
too, will
in
out
days. The KC1 and sucrose
10). The influence of CaCl
5
days.
reproductiveness of
are
that
influence
of five
course
It appears that the
or
amyl-alcohol
n.
get results
to
measured the
we
pollen of flowers that had been
and
9
(Fig.
and
methyl-
variation in the
when
the
on
1, 2, 3, 4
trying
were
Thus,
2
kept in the laboratory for
curves
we
readily.
more
salts,
taken
same
to
we
it
from
day.
use
our
want
will
one
to
be
and
29
Fig.
The
§ 3.
As
on
the
by HQ
ated
of
studying
preceded
the
grains,
a
de
Fig.
into
investigation
we
must
joint
take
20
per
the
influence
into
our
of the
is varied
sugar solution, pu
cent,
and, after measuring the percentage of germin-
by
demonstrates that the
means
region
of the
quinhydrone
where the pu does
11.
depress germination, is comparatively small. In this connexion
the influence of but
salts
Henneman in their
salts
of the medium. It is essential that
the former is determined
11
of
10.
germination.
Jong and
Fig.
not
on
influence
pn
an
germination in
and NaOH
electrode.
by
salts
some
Bungenberg
the alterations in
be
study
by
when
account
Ph
influence
stated
paper,
Fig.
9.
whose
a
few salts may be
behaviour would be
accurately established. Many
worth
investigating
on
account
of
30
their influence
or
too
give rise
colloids,
tolerably
successive
two
Lathyrus
odoratus
that is
have
been
either
low
too
well
in
either
there
season,
(cf. Fig.
Fig.
forced into
the alkali
to
cations;
and
to
be
a
13). In 1938
marked
we
were
12
13.
1939, however,
be muddled. It is
appeared
12
the existence
accepting
in
experimenting
Though results could be reproduced
Fig.
peared
pn
a
we
summers
pollen.
difference between both years
almost
to
high.
During
with
on
of
a
lyotropic series for
the sequence of the salts
impossible
to
find
a
ap-
relationship between
the sequence of the univalent cations and the ionic spectrum of any
known
is
far
less
such
appreciable
relative
there
are
cations,
1939.
differ.
amount
potential
Lathyrus
too,
and
exhibit
When,
for
this
factors that
was
not
example,
will
different
be-
15), though the difference
salts.
might be responsible
grown in the
the ratio of the electrolytes
Possibly,
of Ca,
14
than that found in alkali
many
variation.
a
in 1938 and
brane did
bivalent
1938 and 1939 (cf. Fig.
Naturally,
for
The
colloid.
haviours in
there is
a
same
in the
spots
mem-
difference in
the
greatly influence the action of the
31
alkali
part.
cations
since
antagonism of the ions is playing
The action exerted
the other hand the
by
the anions
solutions
so
also
will
be
an
active
different.
On
activity of the alkaline earth cations will be pretty
(It might be well worth while
constant.
salt
the
as
to
to
study germination
in different
grow Lathyrus
of the
in different
pollen
surroundings.)
This is
borne
Co(NH ) C1
3
AgNOo
6
3
14.
Fig.
15.
by the fact that in both
out
hexol-nitrate
(luteoClg),
correspond
Fig.
to
certain
a
When
amines
act
as
salts.
curves
This is
lengthening the carbon chains,
depression
There
—
exists
and CaCl
2
.
will
occurs
Traubean rule
a
When
1
be lower,
:
strong
keeping
—
constant
by adding CaCl
2
(Fig. 21).
we
16
the
for
curves
ic), T1N0
and
3
and
(Fig.
17). On
18 and
demonstrated
19).
the
The
by Fig.
20.
the concentrations in which
though the ratio governed by the
will
not
antagonism
germination occurring here),
years
page
(Fig.
extent
other hand, the anions show different
hydrochloric
(cf.
can
be attained.
between
50 m.eq.
the
univalent
LiCl, NaCl
make the pollen
or
salts
KC1
(no
grains germinate
32
Fig.
16.
Fig.
17.
Fig.
18.
Fig.
19.
33
From the
experiments under review it would appear that the
of
Lathyrus is
base
not
in every respect
a
suitable
observations of the influence of
our
plasmic membrane. Especially
object
upon
electrolytes
the great
to
the proto-
on
for
sensitivity
pollen
which
changes
in
the pu and the variations in the behaviour towards certain electro-
Fig.
lytes
are
considered in
already
towards
However,
disadvantageous.
that may be
have
Fig.
20.
pointed
alkali
to
cations
the
there
are
the marked
the fact
and salts
that
with
variations in
different
between CaCl
antagonism
§ 4. The influence
of
quite
a
few results
light of the membrane theory.
plained from the antagonism of the ions. We
to
21.
2
may
be
furthermore
and alkali
substances
organic
anions
can
We
the behaviour
ex-
point
salts.
upon
germin-
ation.
In
1938
we
have measured but
did with the salts,
In
these
we
used
compounds,
too,
there
Fig.
few alcohols
a
a constant
is
20
a
per
cent
(Fig.
sugar
22). As
we
concentration.
difference between
Lathyrus
22.
3
34
used in
1938 and that employed in
but these differences refer
and
not
the
Traubean rule.
to
working
the sequence. In
with
alcohols,
in
the
This
1939
did
exception
slightly
a
we
not
of
As
which
importance of the
butyl-alcohols.
have
their influence
with
much
on
24),
confirming
year; all
previous
found
were
to
be
23.
the
influence
act
in
a
molecular
one
of
(Fig.
structure
is
25.
organic
somewhat
substances
higher
concen-
A vivid
24).
furnished
picture
by Fig.
another the behaviour of various
semblance,
And,
germination.
act
in
approach
concentration
each
naturally,
comparatively
26). The action of phenylethyl-alcohol
lower
influence
the
than
that
of
other
takes
n.
whose
closely
phenylethyl-
low
25
isomeric
Tertiary butyl-alcohol and isopropyl-alcohol,
phenylpropyl-alcohol
what
an
in
Fig.
concluded from
compares
23 and
22,
higher concentration than in 1939.
than that of the normal alcohols
formulae
(Fig.
found
methyl-alcohol,
upon soap coacervates, isoalcohols
of the
(cf. Fig.
the absolute concentration
24.
may be
tration
to
occur
Fig.
Fig.
1939
exclusively
in
and
concentration
place
in
a
amyl-alcohol;
some-
it
is
35
reminiscent
same
takes
is
felt;
felt
coacervates
of
activity
by
n.
on
the
practically
phenylethyl-alcohol
amyl-alcohol.
only in fairly high concentration that ether (Fig. 26)
exerts
germination. The influence of dipropyl-ether
in much lower concentration.
Ketones
(Fig.
centrations
found
acetophenone
ethyl-ketone
Lathyrus
ketone,
too;
27),
too,
with soap
experiments
of
here,
oleate
the
that shown
over
depressing action
is
In
hexyl-alcohol.
precedence
It is
a
of
influence
<
Fig.
27.
behave
oleate
—
We
we
as
coacervates.
in
pollen.
its
26.
The
might
apparently
<
expect
sequence
coacervates
methylbutyl-ketone
acetone
<
Fig.
—
occurs
lies
between
that
also in the
of
from
active
the
con-
dipropyl-ketone
diethyl-ketone
measured the influence of
activity
of
<
<
methyl-
germination
methylpropyl-
diethyl-ketone
and
methylethyl-ketone.
Carbamates
(Fig.
(urethanes)
28); this again
oleate
in the
coacervates.
bring
may be
out
once
compared
more
the Traubean rule
directly with the action
Glycol, glycerine and propylene glycol
neighbourhood
of
sucrose
(Fig.
23 and
29).
are
on
found
36
We
saw
is
chains
carbon
atoms
coacervates,
exert
the
a
depressing
we are
stands
This
on
trying
For
<
picture
and
the influence
fact
agents
while
83).
action
to
on
with
act
as
Fig.
28.
Fig.
29.
germination.
oleate
might well
For
each
that,
in
others
coacervates
we
that (in oleate coacervates)
opening
will
be
<
isobutyl-alcohol
<
propylcarbamate
influence,
(This
having
and
condensing actions.
shown this
<
<
series
been
compare
homologous
advantage of showing up clearly
isopropyl-alcohol.
of the
We
another.
one
find the compound
has the
propyl-alcohol
ether
the number of
to
opening
series
Diethyl-ketone
n.
likened
be
in the
the border-line between
procedure
ences.
concentration,
The main difference lies
compounds
some
organic substances in homologous
lower
which may
increasing,
homologous
series
a
shrinking action, whereas all of these organic non-electrolytes
a
exert
readily felt in
coacervates.
soap
on
that the influence of
more
sec.
tert.
gives
the differ-
sequence:
butyl-alcohol
butyl-alcohol
but
reconstructed
an
<
<
imperfect
from
17,
33
37
The influence
germination of Lathyrus pollen is practically
the
on
identical:
Propyl-carbamate
butyl-alcohol
sec.
alcohol
<
diethyl-ketone
<
ether
<
<
the result is
data of various
over, in
preparations of oleate
the
germination
occur
§ 5.
(ether
and
of
Stability
the
influence of many
that
carbamate)
distance between the first and
.85. Hence, the few variations
of minor
are
importance.
in distilled
remain
on
When establishing
the
germination of the grains it
water.
the
intact
in
certain
concentrations.
bursting of the grains is easily investigated with the aid of
extinctometer,
the
as
of
disintegration
cloudiness
the
the
into the
placed
now
In
thermopiles.
source,
the other
receive
a
contains
different
one
to
zero
of the
of the
by
be
We
cloudiness
we
the
are
containing graded
As
means
sugar.
fast
of
The
then both
as
a
stability
preparing
sugar
rheostat
that is
measure
pipette,
amounts
of
two
cm
will
thermopiles
pollen
are
be
brought
circuit of
The
rheostat is
light intensity.
sucrose
too
on
(10
cm
cm
introduce into
3
can
in the
placed
germination
grains (Fig. 30).
especially
hours
light
while
light dispersed and absorbed
series of
a
flasks
constructed
piration of these
of the
we
2
water
of resistance will be indicative
concentrations
possible
two
galvanometer
light between
side of the
dispersed and absorbed.
light
Erlenmeyer
wooden tray,
the
a
that the
way
on
either
the
of
light, in other words the galvanometer
comparing the influence of
that exerted
by
of
thermopiles. The
of
on
A
of
source
of these cups contains
read in percentage of the total
are
flask).
of
a
The
Moll’s
with
consists
position of the galvanometer
means
amounts
graded in such
may
one
pollen suspension,
a
amount
will be deflected. The
back
circuit and
front of the latter,
If
considerably
extinctometer
oppositely connected.
are
common
placed.
are
cups
increases
This
grains.
thermopiles whose circuits
is
the
grains.
compounds
these
by different
coacervates
mutually compared. More-
are
logarithmic
bursts
Lathyrus pollen
appears
<
butyl-
tert.
established
were
conspicuously fine. In oleate
the last substance of the series is but
that
<
isopropyl-alcohol.
Considering the fact that these data
observers,
isobutyl-alcohol
<
propyl-alcohol
n.
3
3
for
the
in each
the
Erlenmeyer
Erlenmeyer flasks
in
shaken for
this
with
measure
Erlenmeyer flasks
suspension
then
To
20
2
purpose.
the cloudiness of the
per
hours
On
cent,
in
the
a
ex-
liquid in each of
Erlenmeyer flasks is quickly measured with the extinctometer.
The
values shown (percentage
of absorbed light) are
against the sugar concentrations (Fig. 30). It appears that.
resistance
plotted
38
Fig.
in low sugar
centual
concentration,
that
cloudiness)
sugar concentration
such
to
In
an
order
sucrose
to
and 5
3
Once the series is
suspension
for
20
2
per
in
hours.
cent,
20
As
no
grains
the influence
salt solution
prepared,
per
no
cent,
we
(high
in.
sets
In
surroundings increases
of
add all
salts,
5
into the
cm
3
40 per
along the line
2
cm
are
demonstrated in
Fig.
30,
the
concentrations,
cent,
Erlenmeyer flasks.
which the flasks
sugar, upon
sugar. In certain salt
per-
high
longer burst.
passed
are
rapidly
so
germination
the osmotic value of the
measure
cm
burst
grains
practically
that the
extent
the
30.
grains
will
3
pollen
shaken
burst
in
however, the grains
remain intact.
According
ing
to
sequence
their
(Fig.
protective action, alkali salts rank in the follow-
31):
Cs
>
Rb
>
Na,
Fig.
31.
K
> Li.
39
The sequence,
quite
with respect
to
depression of the germination, is
a
different:
Na > K
In alkaline
from that
earth metals
on
the
>
of the
Fig.
germination
this
while the
is
sequence
Ba
>
following action
Mg
Mg
About the influence of T1N0
3,
hexol-nitrate
reported
(Fig.
upon
33).
the
>
Rb.
grains
on
(Fig.
varies
germination
32).
33-
shown:
> Sr
the
on
Sr >
Cs
32.
Fig-
In
>
the influence
also,
bursting
Li
Ca,
stability of the grain exists:
> Ba
AgNOa
stability,
>
> Ca.
,
Co(NH )6 CI 3 (luteoCl 3 ) and
nothing
3
of
importance
can
be
40
Fig.
34-
Fig.
35-
Fig.
36.
41
Fig-
37-
Fig.
38.
Fig-
39
42
Normal alcohols
the
be found
and
great and
too
Contrary
stability
show
and
§
the
6.
of
sequences
comparison
ethyl-
(Fig.
34).
germination
and
have drafted the
we
only methyl- and
Isoalcohols
(Fig. 35)
alcohols.
The
act
curves
then cloudiness
increases.
Butyl-alcohols
picture of the development of this minimum
Respiration.
first
at
centrations,
(Fig-
measuring the influence of
results
respiration,
upon
salt,
a
not
were
at
in
divers
con-
tangible
once
37)-
To
respiration
measure
the
prepare
6
taining
vessels
usual series
3
of the
the
of the
cm
we
of
Warburg’s
use
test
tubes
Of this,
liquid.
central
in
20
per
cent,
in the
the results
Fig.
demonstrated in
salt concentration
lined up
exerts
a
cm
parallel
toxic
to
action
3
the
the
x
is
usual
37
KOH is
manner
fairly
(43).
easy
but the
1
3
cm
vessel.
Ultimately,
explain.
to
the
grains burst, consequently
axis. In
the
passed
pipetted into each
were
con-
is introduced into
pipette.
a
sucrose
We
(44).
Immediately before the experiment,
cups.
Respiration is then measured
In low
1
apparatus
§ 2), each of which
(cf.
Warburg apparatus with
pollen suspension
are
small
too
maximum.
subsequent
When
into
as
but
fine
a
us
concentration than normal
minimum,
a
here, the
standard of
a
to
in osmotic concentration.
act
higher
36) give
shown
From this it would appear that
curve.
slightly
(Fig.
As
concur.
sugar
be
electrolytes,
to
ethyl-alcohol
in
though on
propyl- and ethyl-alcohol will
the other hand the distance between
on
will
methyl-alcohol
cane
follow the Traubean rule,
generally
hand the distance between
one
curves
high salt concentration the salt
grains
remaining
intact,
we
obtain
straight lines. The inclination will be indicative of the salt’s toxicity.
To
we
compare with
plot
the
one
amount
another the influence of the salts
of 0
taken up in
2
rithm of the salt concentration. As
sugar,
the blank will
But little
It
appears
and
AgN0
that
17) than
grains
only
to
in
concentration
This
is
3
be
is needed
to
on
respiration,
against the loga-
grains burst in
20
per
cent,
keep the pollen grains intact (Fig. 33).
AgNO a is much
more
toxic
to
germination (Fig.
16
respiration (Fig. 38). Alkali salts stabilize the pollen
high
they
concentration
exert
the sequence of
a
(Fig.
39).
(Fig.
31).
depressing action
activity
Of the alkaline earth cations
metals
the
minutes
fairly low.
K
alkali
too
on
In
almost
the
same
respiration (Fig. 38).
found:
> Na
> Li.
Mg behaves
The other
salts
in
some
present
measure
a
very
like the
different
43
the
Here,
picture.
centration
grains remain intact in
while
region,
the
fairly extensive
a
action
depressing
respiration
on
con-
only
concentration. As the curves are measured
a higher
days, the peaks must not be compared. Only from the
manifests itself in
different
on
position of the peaks
that
can
similar
closely
are
ascertain that Ba and Sr
we
in
while
strength,
Ca
actions
exert
exerts
weaker in-
a
fluence.
CHAPTER
INFLUENCE OF
THE
VARIOUS COMPOUNDS
BAKERS’
ON
§
III.
YEAST.
Introductory.
1.
Before
like
entering upon
examine
to
These
on
(4,
of their
It is
When
this
should increase.
occurs
more
share,
influence of
known
on
the
C0 2
that
saponin
lipoids
be
can
fermentation along
on
precipitated by
he
than
found confirmed;
cells
that
not
were
yeast
cells
Kolloid aufzufassen ist, und
so mils
Lipoid,
sen
die Salze
zwar
genau
Plasmahaut verandern”. Lecithin
S0
>
Cl
N0
>
4
>
3
From
same
this
order
he
this
than
properties. In this
while
depression is
lipoids;
influence
the
to
salts,
on
the
greater
of
salts
Hauptsache
als
hydrophiles
salts
following
this series:
stimulating action will decrease in
might
NaCl
In a
observed.
apart from
deduct that
exerts
a
structure
the
on
than
influence metabolism; “Die
den
Stoffaustausch ein,
da
to
the
has
to
oblong form
the
BaCl2
a
on
stimulating
considerably
Boas, does but influence
membrane,
having
the
some
mem-
through which the influence
noticeable.
In
alkali
cations
in alkaline earth cations.
Lipoidhaut
sie
depressing action
elliptical
and
the
greater
so).
The action of saponin excites
for electrolytes
becomes
und
not
strongly
Saponin, according
action
a
2
mixture of saponin
their
has
CaCl
compounds individually
structure.
permeability
so
the
these
with greater clarity
stark sein diirfte,
der
some
die Plasmahaut
bekannten lyotropen Reihen die
precipitated by
one
will
rein
is
in
den
the cells change from
plasmic
protoplasmic
observed
that
and
vacuoles disappear.
smaller
brane
case
wohl
nach
NaCl, which, however, is
each
of
SCN.
theory
mixture of saponin
fermentation while
the
>
concludes
(from
stimulating action
A
J
means
treated with
Also
treated.
(for example NaCl) stimulate fermentation. Moreover “da
als
the in-
whole, the authors’
inside the protoplasmic membrane, permeability
This, indeed,
saponin liberate
not
others.
by
with
dealing
should
we
out
findings.
established the
5)
gravimetrical lines.
saponin.
experiments,
own
carried
the membrane. We do
interpretation
Boas
our
experiments
important because they, too, are
are
fluence
review of
a
number of
a
wirkt
mbglicherweise
demnach
nicht
zu
this
will
on
be
Naturally, this
regulatorisch
auf
jeder Zeit gleich
sind damit mancherlei Regulationen des Stoffaustausches
physikalisch-chemisch
gegeben.”
44
A12(S0 4)3 exercises
of A1
a
and
(S0 4 )3
2
depressing influence
a
saponin
and
mixture of saponin, NaCl
of
saponin
and
HC1
has
a
fermentation. In
on
C0 2 is liberated
more
2
discovered
(58)
that
fermentation of glucose. This
he
terin. When
mentation in
increased
washed
distilled
this
on
to
the
Hermann
specific action.
a
though
not
does,
In sols
put
of
In
salts.
down
we
shall
As
we
have
time.
a
experiments
C0
2
this should
we
do
only
room
get
better
a
rate
of
to
to
for
long
too
during
one
in
a
lecithin
and
obtain any
not
cholesterin
with
precipitate
must
yeast
upon
This
substances
numerous
into
insight
measure
a
we
the
be
would
fer-
upon
of the
structure
.
the influence of
shall
have
to
we
—
alcohol
are
to
—
carry
shall, therefore,
We
are
suitable
not
considerable
a
for
or
a
method
series
at
the
the remainder
out
number of
a
have
to
measure
error.
2
quantitative research,
The
research.
Many
other
fermentation flask and
disadvantage. On top of
which allows
research.
methods
a
gauge
methods alike
planned with
a
view
measuring
Kluyver-Van
to
define the
the
amount
yeast
for
to
measuring
marked
are
the yeast mixture there is
respiration of the yeast
The
use
liberated. Now, the apparatus consists
Gravimetric and volumetric
common
we
gravimetric method leaves
2
for
compounds
find
fairly comprehensive
time if
day.
preliminary
of two separate parts;
the C0
able
an
fermentation.
measure
measuring the volume of CO,
this
be referred
not
the membrane cholesterin.
upon
concentrations,
us
Einhorn tubes
of air
From
depression
liberated.
them
a
cholesterin.
specific
speak in favour of their experiments.
Defining either the final product
glucose takes
the
views
own
used ill-defined
not
undertaken
many different
which will allow
of
exert
permeability.
the
Measuring
2.
had
toxin
a
layer of the protoplasm.
outer
in
we
fer-
found the
for
permeability of the cell, cholesterin plays
the influence of
measure
mentation,
our
must
the influence of saponin
action
an
to
have
the increased
explain
If
to
the
choles-
specific part.)
a
opinion
he
is also opposed by
cholesterin
lecithin
purified
our
not
the
the whole,
on
stimulate
depressed by
consequence of the membrane theory,
influenced by cholesterin.
co-zymase
that
In
That the authors
sols
mixture
direct
a
yeast juice is
concludes
be
can
with petroleum ether
yeast
fermentation. (According
as
a
addition of cholesterin.
preparations
unaffected, though sensitivity
stimulating action of
important,
§
the
As
insulin
some
stimulating action
water
considerably.
fermentation of
The
than
less injurious action
NaCl.
Obviously, fermentationcan also be depressedby
Hermann
mixture
a
in A1 (S0 4) 3 alone. Also,
than
to
Iterson
amount
activity
take
an
has
of sugar; it
varying
is
not
by
layer
active part
apparatus
under
a
in
been
suit-
conditions.
45
Warburg’s apparatus
for
which
As
none
will
it
be
accommodate
cannot
this
reason
method
(80)
of these methods
follow
to
necessary
can
neither
are
be
suitable for
rest
will get
be
is filled with
Our first apparatus
partly put inside
differences
in
fermentation
thermostat.
some
2.
a
to
which
are
branching off
new
at
the
for
paraffin
the side and
fermentation
This
gave
necessitating
could
be
(cf. Fig. 40)
to
the
construction
an
impossible
to
obtain,
due
error
immersed
completely
only
was
rise
type
from them,
of
in
to
a
the
during
figures
reproduced.
Fig.
40.
Fig.
two
factors
factor
is
that
of the
yeast will settle
The first
long
In this
tube which
have used fermentation tubes of this
supersaturation
the
experiment,
glass.
temperature
We
tube
measure
time, but found it
There
little
thermostat.
a
tube
that could be
1.
a
measuring
a
this
oil; if C0 2 is liberated,
paraffin
pushed through
collected in
a
series
used.
fresh line of conduct.
a
method the yeast mixture is enclosed within
the
comprehensive
a
detrimental
to
results:
with C02 will
occur.
during the experiment.
of
paired by introducing
are
yeast suspension
41.
a
This evil might be rephase into the liquid phase of the
great consequence.
gaseous
6
4
tube.
This
have tried
we
saturation with C0
with little
it
set
in
effect in
to
did
2
rings made of the
when
pipe
a.
A
of Sambucus
of error
sources
used
by
has
us,
fermentation tube. A
several
(a)
tube
is
2%
is
which has
fermentation
fitting
b.
in
with
provided
long,
cm
tube is
can
be
good grip
closed
with
cm
3
In the
).
diameter
towards
pushed
moderately
a
a
50
thick
a
mm),
tube, abt.
tube.
glass
perforated,
a
1.7
the exterior.
rubber
the
on
across
centre
paraffined
The
cork
the rubber tube.
A little
C02
any
glass eyelet. To stir the yeast suspension without
a
escaping, the stirrer wire is conducted through
glass tube which
by
means
of
down than
is
cement.
To
three
This
80
which
mersed
by
in
lities,
a
(Fig.
toluol
be
placed
taneous
annular
glasses
Stirrer
copper
is
a
is
vessel
that
As
as
collect the
to
a
synchronous
an
eccentric.
The influence
demonstrated in
Supersaturation
by
at
im-
38.3° C.
means
at
of
two
the time
possible
a
can
simul-
Around the thermostat
which
we
put
the
an
measuring
over
the
thermostat
there
is
a
be shifted up and down. The stirrers of the
are
43.
with C0
2
onto this ring. The latter is moved
(driving also the thermostat stirrer) and
hooked
motor
exerted
Fig.
of abt.
completely
constant
heated
it.
onto
paraffin.
Suspended
can
fermentation tubes
by
on
of the fer-
fused
capacity
a
18 fermentation tubes
constructed
installation.
ring that
held
The apparatus is
many
little further
little hook.
are
the thermostat which makes
is
are
tubes
are
a
having
the fermentation
a
centre
(d)
with
provided
copper
examination of many media.
table
in the
The temperature is
42).
inside
the tube
keep
wire
regulator.
Argand burners.
narrow
tube branches off the
glass protuberances
The end of the stirrer
Thermostat.
This tube reaches
where the other little
point
tube.
mentation tube,
a
glued inside the cork of the fermentation tube
Kothinsky
the
fermentation
d.
removed when
glass tube (c) is closed by fusing the bottom. A
piece of lead is then introduced, after which the top is fused and
Stirrer.
provided with
c.
second
component parts:
(internal
branched off
through which the paraffin
The top
are
did
nor
long, glass tube is closed straight
by fusing its bottom (Fig. 41, capacity
capillary
provided
nigra;
in the fermentation tube is stirred.
suspension
The apparatus,
No super-
rods
glass
employed. Nevertheless, the
objection would still be felt. Both
the yeast
used
we
marrow
were
stems
different ways.
two
when
occur
by
stirring
At
A
will
on
the
the stirrer
set
in
liberation
of
installation is
immediately
upon
C0
2
is
stopped.
this and the
47
Fig.
42.
48
curve
will flatten
opposed
out.
When stirring
and the line will
Fig.
To
demonstrate
with
experiment
The
the
anew
method
different
following mixtures
of
amounts
are
% yeast suspension
10
water
now
is added. The stirrer
that the
branch-tube
paraffin.
A
of
attained.
we
shall
review
an
yeast.
5
cm
2
3
33
8
„
»
d
c
e
3
3
cm
2
2
2
3J
10
it
„
cm
3
I
cm
2
35
12
»
3
33
„
poured into the fermentation tube and paraffin
is
then
placed into the fermentation tube taking
glass tube is filled with paraffin
the fermentation
little
is
suspension is added immediately before the experiment.
The mixture is
care
3
53
3
The yeast
procedure
b
cm
2
curve
prepared:
.
30 % glucose
original
43-
a
io
(B) supersaturation will be
steepen until the
note
tube
we
is
put
inside
placed
a
25
3
cm
the
up
measuring
glass
is made of the time. We will then
Fig.
44.
to
the cork.
to
Now
under
the
collect
the
thermostat;
measure
at
regular
49
intervals,
five
say every
minutes, the
As the
do
amount
of
indicate
paraffin collected.
the
correct volume,
measuring glasses
exactly
they will have to be graded in advance. These readings we corrected,
using the data in
This
of
This method
the
and
will?not
Let
be
a
collected in
is
rate
this
be
to
the
mically,
t
the
as
between
=
C
—
a
log. G,
Ca
G
this
need
io
=
K
=
this
3
3
cm
and
3
20
to
decreases.
G,
we
must
The
denotes the
cm
) divided by
a
and
paraffin.
amount
(t ) required
(G)
3
t-2o-io
of
paraffin.
paraffin
t 20-10-
collect
If
we
plot,
3
a
cm
want
or
.
two
and
x
time
y
to
in the
relation of
simple
amount
time
than
one
does
to
find
not
Here,
relationship.
the
of yeast always needs
take in
would
be
an
equal volume
expected
from
relationship between the amounts
of yeast
To
(46, 79).
measure
trolytes
series
on
.
the influence of elec-
fermentation,
of
we
experiments
concentration
of the
carry
out
alternating
electrolyte.
Fig.
45-
4
,
to
logarith-
that:
.
a
the
of
cm
by the
It appears
that
yeast
lapse
ta
cm
a
x.
smallest
a
of
stand
respiration
more
When
paraffin
given volume of Co2 These
now
and
In
a
a
would result
different
liberate
times
it
amounts
a
ordinary
Thus,
.
10
10
of yeast
(Fig. 45).
ta
different
the
v
that the time
amount
G
against
a
X
From
O
collect
be
lapse of time (i.e.
a
y
to
represented
relationship
t
log.
is
V20-10
expected
increase
learn
reckon with
paraffin.
the results.
to
time needed
the
a
average
It
the
series with
passed between the reception of
The
we are to
—
collected each minute will
average
time
a
—
the action
result will be found.
we
with paraffin
pre-saturated with C0 2
figures. Here, the solubility of C0 2 in paraffin
detrimental
denote
t
2
in
44).
defining
series
identical
at
dissolving
of blanks
Fig.
same
error, since
to an
C0
some
series
two
another
arrive
will
of
the
days, practically
the numbers found
(cf.
When
reproduce.
might give rise
eventuality
comparing
readily
different
on
upon which
possession,
the time in minutes
method will
salt
a
our
plotted against
were
not
50
The
following
different
14
KCl
KCl
I
3.5
n
.35
n
.035
KCl
—
30 %
1
After
4
3
4
3
—
—
2
4
5
2
2
2
add, with
we
shaking
a
6
5
2
_
—
8
7
10
9
II
12
13
—
tube Nr.
put
great
too
a
2
8
6
2
2
—
twice,
2
4
—
—
—
—
7
2
4
2
2
2
cm
—
6
-
—
—
—
2
—
—
2
6
2
6
2
2
2
2
2
2
cent,
7 per
the mixture
is
make the fermentation tubes face
10
another
one
tubes
the
yeast suspens-
poured into the
way. As
not
we
to
group A
full minute. Half
will be read,
do
moving ring,
The first
the full
minutes,
the half minutes. Also
to
of
manner
we
(Fig. 46).
timed
are
others
reading
same
side of the
one
2
8
placed in the thermostat in the
on
6
6
3
is treated in the
charge
etc.
a
so
takes
place
at
pleted in
that all the
of
numbers
the
bers
Fig.
a
average
rate
amount
of
total
KCI
rate
of
curve
upon
describing
fermentation
of
10
2 or
in
this
3
of
20
cm
3
readings
chart
com-
adjusting
the
potential
the
glasses,
plotted against
some
of the KCI
Here,
this
particular
This
the
num-
in the above
is
take the
we
the
average
dealing
established
with
a
for each
relationship of these numbers with the
is
defined.
(In this
case
the
corresponding with the numbers 4
experiment.)
way
are
(Fig. 48). Here,
to.
representing
minute when
one
glucose alone
blanks is taken,
referred
the influence of KCI
paraffin).
which the
KCI concentration
the
47
reproduced.
are
(v-2o,
The percentages found in this
the
Fig.
correspond with those
collected in
of fermentation in
of
to
measuring
found will be
In
the
chart.
for
paraffin
quantity
average
and
46.
criterium
After
paraffin
the
time.
curves
As
of
amounts
errors
minutes.
two
the
minute later group B
of the fermentation tubes will be
the
14
I
pipette, 5
the tube
fermentation tube which is
to
introduced into
are
n
To tube
want
example,
6
n
Water
ion.
(cm ),
2
6
.0035
Glucose
for
tubes:
test
Numbers
KCl
3
mixtures
plotted
too,
against the log. of
the numbers
are
those
51
Fig.
47-
Fig.
48.
Fig.
49.
52
Fig.
50.
Fig.
51.
Fig.
52.
53
Fig.
53-
Fig.
54-
Fig.
55-
54
The
§ 3.
The
influence
influence
measured in the
of
of
salts
same
fermentation.
on
various
salts
manner
fermentation
on
KC1. Alkali salts all
as
of
glucose
exert
is
stimul-
a
This stimulating
influence of the univalent cations
ating action.
(Fig. 49) diminishes in the series:
NH
> Na
>K
4
the
earth cations
Ca
does
not
following
act
Li, being
>
stimulating
in
for
the
Sr
low
as
salts
heavy metals, in
action
pressing
(Fig. 52)
measured
a
study of
do
not
in
have
the other
of
a
n.
.04
mentation
Other
salt
a
curve
curve
has
gets lined up
Boas
increases
salts
show
a
were
these
the same,
high
they
as
a
this
been
From
action.
stimulating
hydrogen ions
they
becomes
depression
we
concen-
have
must
are
con-
exceed
not
influence
fer-
on
increases.
remarkable
(Fig. 53, 54 and 55).
curves
low
comparatively
no
influence,
parallel
a
concentration.
the
parts of the
(4,
x
at
much
higher
This
are
concen-
which
at
varies for each
axis,
curves
the
5)
until
second time. This level,
to
salt.
the
Also
different.
action
stimulating
of
the
anions
the series:
employed
similar
Br
in these
behaviour
<
<
3
While the influence of Th
addition of NaCl
(Fig.
Cl <
S04
.
measurings (Fig. 56).
(Fig.
no
the
de-
a
of
toxicity
times
.
increasingly depressing
CNS <
K
the
2
acid reaction;
very
drop
observed
in
a
very
will
the initial and last
As
50);
definite level is reached. A further increase of the
concentration
tration, the
10.000
exerts
influence upon fermentation until
an
in
occurs
U0
much
as
the same,
Below p
H 3,
has
salts exhibit
Depression
already
The
$1).
Th <
<
For
salts.
concentration
continues until
earlier,
literature it appears that the
depressing
TINO.j
as
(Fig.
Hg
<
observe
we
(N0 3)2 has
Be
spicuous.
to
been examined in
not
high concentration.
abt.
Ag
alkali
existing
the
exert
concentration that is
referred
(Fig.
observe
the series;
Cu <
RbCl and CsCl
a
we
> Ca.
> Ba
fermentation
on
in
increases
tration
Mg
case
action:
stimulating
>
that of the salts
as
lyotropic series. In alkaline
somewhat weaker
low concentration. In this
sequence
A group of the
normal
a
action is
Cl <
salts,
too,
so
4
.
(NOs )4 practically
58),
Mn
57):
we
do
does
observe
a
not
change
clearly
on
defined
55
Fig.
56.
Fig.
58.
Fig.
60.
Fig-
Fig.
57-
59-
Fig.
61.
56
between Ni
antagonism
of NaCl is
The
§ 4.
On
that,
(.53
needed
influence
fermentation
to
a
of
the
that, here,
compounds
on
influence
state
the
cause
organic
compounds.
normal alcohols
we
of alcohols
on
yeast and
acetone
> 5.0
According
occurs
in
to
juice.
„
.23
,,
62.
Fig.
63.
their observations, in these concentrations
by
to
Fig. 63 where
a
us
that there is little
direct
action
the action of
The volume of
yeast
doubt that
some
cells
so
we must
closely
attribute
fermentation.
on
structure
becomes
clearly manifest
phenols is demonstrated.
in
salt
(4) observes that the yeast cells lose
mixture of
precipitation
juice. These concentrations approximate
yeast
influence
an
They
further
,,
.54
The influence of the molecular
assume
no
„
1.3
isoamyl-alcohol
the values found
Boas
of these
(98) established
yeast
isobutyl-alcohol
Fig.
a
action
60)
it is
mol.
3.5
propyl-alcohol
n.
in
However,
occurs:
ethyl-alcohol
§ 5.
an
Warburg and Wiesel
Methyl-alcohol
in
with
(Fig.
action
an
exert
the Traubean rule.
concerned
are
permeability.
amount
shift.
to
curve
having found the following concentrations in which
fermentation
this
59). A fair
3
to
certain extent, follows
improbable
the
(N0 )2 and NaCl (Fig.
n)
solutions.
their natural
appearance
saponin and NaCl. Vacuoles disappear and the cells
oblong shape. This he puts down
action of saponin, gets
an
opportunity
to
to
the salt that,
by the
invade the cell. We observe
57
identical
Th
changes
(N0 3 )4
invasion
is
It
the
the
on
(N0 3 )2
and U0
2
addition
of
Here, again,
.
CuCl
AgNO s
2,
we
HgCl
,
2,
may be faced with
an
of the cations.
also
other
osmotic
interesting
very
salts
in
high
phenomenon,
ascertain
to
concentration
or
An
not.
whether
should
answer
the influence of
be
regarded
these
to
as
an
questions
is
by measuring the volume of yeast cells in different salt
furnished
concentrations.
For
20
these
3
cm
measurings
These tubes
tubes with a capacity of abt.
use little
tapered towards the bottom to a graded
we
are
.
piece of abt. 3
vate
10
cm
3
cent,
per
In
solutions
2
then be necessary
this
we
some
ten
deposits,
It
U0
2
would
cells
(La(N0 )3
to measure
now
that
appear
exceptions
to
this
and
Th
(N0 )4
on
3
indeed
be
so
as
to
arrive
practically
are
.
2,
Pb
(N0 3)
2,
uniformly. It will
immediately.
). The long and the short axes
.1 n
the five
all
the volume. As
at
salts
completely:
It is
a
in
blank.
of the salts diminish the
and
1
(Fig.
n
AgN0 3
to
64
and
that, in low
probable that
the cells. With respect
found
in
becomes noticeable.
being
for
account
influence
an
on
,
CuCL,
these
salts
65).
concen-
HgCL 2
,
exert
a
other salts, the second
osmotic
influence
65
few
we
1) We
are
apparatus
on
upon
measuras
volume.
have sketched
examples
of
phenomenon.
again measured
are
volThis
the fact
fermentation has been
their
the
fermentation
(All salts whose influence
a
3,
settle
of concentration
region
would
Fig.
3
not
of the fermentation lines
where the influence upon
ed,
CuCl
Ce(N0 )
,
x
(6)
between
fermentation
depress
3
very
this
3
cm
the volume of the yeast
the volume of yeast cells
measured
are
direct, toxic action
ume
coacer-
solution and 8
compare the numbers obtained with
(N0 )2
curve
will
to measure
salt
cm
days
two
(N0 3) 2) yeast does
volume in concentrations
tration,
After
Box’s apparatus
use
we
The sole
7
pass
3
Th (N0 3)4 and U0
of
often used
are
3
we
measured.
few salt
a
suspension.
yeast
is
deposite
For
tubes
(similar
volumes). Into the tubes
of
to
In
but
salts; those
Fig.
64.
extremely grateful to Professor Box for lending us his measuring
and
for
demonstrating
to
us
its
use.
%
58
not
charted
§ 6.
are
found in
Respiration
will have
to
65.
Fig.
66.
Fig.
67.
exactly the
same
region of concentration.)
of yeast.
When the volume of
get smaller.
Fig.
a
Respiration,
yeast cell
diminishes,
which is
regarded
follow this decrease. We, for us,
as
are
its
a
surface,
surface
(Ni(N0 3) 2 behaves
respiration.
like
this)
will
exert
will
concerned with the
problem whether salts that bring down fermentation
level
too,
catalysis,
a
to
similar
a
definite
action
on
59
In
number of
a
and
glucose
vessels
8
3
transferred from the
ration of the mixtures
It would now
Respiration
curve
is
No
than
more
addition
of
is
But CuCl2 has
a
fermentation and
centration
tube 5
test
Subsequently,
cm
2
cm
3
3
3
respi-
3
different from that exerted
2
ferment-
on
region where the volume
in the
depression of respiration is
a
do
observe
we
2
NaCl
or
cm
(Fig. 66) that the action exerted by Ni(N0 )
Ni(N0 )
KQ
pipette.
2
each vessel upon which
to
only depressed
3
adding
into each
of
the Warburg
as
decline.
to
seen
a
soon
be measured.
materially
ation.
tubes
As
pass
of
means
test
will
appear
is
respiration
on
we
mixtures
preparing
are
solution.
with KOH,
suspension (2 %) by
yeast
are
we
Ni(N0 3) 2
provided
are
tubes
test
cm
(Fig.
a
to
occur
on
action
the
when
67).
powerful action; here, the
volume
seen
stimulating
found
are
of
curves
in
practically
the
respiration,
same
con-
region.
CHAPTER
IV.
GENERAL CONCLUSION.
§
of
In
the
substance
a
give rise
to
can
can
be
act
which
the
structure
derived.
upon a cell in
two
different
manners:
change in the membrane.
a
The influence upon
2.
from
experiments
membrane
general,
It may
1.
the
Selecting
1.
an
internal system
of the cell
manifest
may
itself.
Though, also in
with
action
I, § 2),
ter
a
an
the
shall
not
these
practical
Lathyrus
pollen is
variations in the
interpretation
the
to
cause
There is
able,
a
use
not
least in the
that
we
a
shall
very suitable
germination of
the
future,
near
of these
variations.
relationship
further
For
Chapto
of these internal systems.
in the solution of
select
our
object.
giains (Fig.
the
Firstly,
9 and
the
present
we
give
It is
experiments
membrane
problem.
the great
10) make such
can
between the weather and
to
fathom
only
point
germination.
relationship between the colour, the specific weight
germination. Pollen, showing
with
at
nature
faced
be
may
difficult. Further research would be needed
the apparent
and
be
grounds
which may be of some;
we
complex and symplex relations (cf.
interpretation of the
correct
on
we
the last-mentioned instance,
upon
difficulty; it is coloured
a
great specific weight, germinates
orange.
Pollen that
germinates readily
60
has
of
yellow colom
a
in
water
the
As
the
is
fluence of
spectrum of
impossible
in
study of
our
an
we
object,
be
This,
only
of the
measure
on
the in-
germination shows
directly likened
results
shall be
we
amount
action
injurious
an
can
for the time
of the
The
weight.
part.
exert
n),
cannot
interpretation
an
not
Fig.
which
biocolloid.
a
does
(cf.
specific
active
an
Now, depression
of the salts
sequence
small
a
play
pH
limited
few salts.
a
but
may
where
region
germination,
and
grains
being
a
the ionic
least, renders
at
obtained.
permitted
to
Only when,
experiment with
to
various
salts, can we eventually arrive at important conclusions.
Hydrochloric amines behave as salts, though the influence of the
readily felt (cf.
carbon chain will be
reversal
L.
as
sequence
of the
Teunissen-Van
the cation
lengthens,
character while the concentration
The sequence, exhibited
of
germination,
brane
upon
same
the
established
derive
to
assumes
of a
more
the reversal of
at
hydrophobe
charge decreases.
by neutral salts with tegard
be
directly likened
some
from
to
to
a
depres-
ionic spectrum
an
the
behaviour of
conclusions
with respect
the salts
the
to
we
mem-
With a view to the stabilizing action of these salts
grains it becomes evident that the influence on the membe
must
germination
salts
pollen
organic substances
the action
we
can
consider
complex
the electric
the
direction
the
upon
on
re
the
condition),
For
at
inside
the
in Lathyrus
hydrophobe
All
of the
part
practically all
between the
of the substances
along
§
two
2
we
seem
shall observe
different lines
bursting of pollen grains give
of the membranes in various
stability
of
curves
germination and those representing
(when reading
be arrived
can
depressing
eventuality
relations
foretell the relationship
the influence
Measurings
of the
act
of normal alcohols.
same
depression
mainly alter
influence
they
when
however,
must
(i.e. they influence symplex relations).
representing
in the
we
internal system.
an
compounds
organic
membrane
on
salts,
electrolytes
(i.e.
factor;
important
an
with alkali
acting
Whereas
membrane
a
who
structure.
the
brane
these
cannot
Nevertheless,
biocolloid.
a
shall be able
(94)
charge of biocolloids with the aid of amines. When
the carbon chain
sion of
Fig. 20). We observe the
Zijp
of
some
to
work
that
this
conduct).
impression
surroundings.
This is
problem much less complex than germination; when the substan-
ces,
whose influence
osmotic
bursting
quantity,
we
phenomenon
we
are
shall
about
have
directly
to
to
to
measure,
ascribe
changes
are
their
not
action
within
the
present in
upon
the
membrane
itself.
What holds
applies
good for the influence of electrolytes
equally
to
the action
on
respiration:
on
germination
the number of salts
61
we
actually
can
of the results.
will burst
alcohol
curves
respiration
for BaCl
blank) burst after
to
almost flatten
serve
the
more
rapidly.
we
On
adding 5
m.
10
in
mol.
longer since the respiration
action
(30
section
from
the
(blank)
68),
in
amyl-alcohol;
20
mol.
m.
Whereas
they
the
Lathyrus pollen
yeast
one
When
we
grains
may
influence of
m.
curve
under
mol.
sugar
cause
the
curve
ob-
we
though this will
set
remain
grains
same
(i.e. the
amyl-alcohol,
n.
amyl-
in
intact
68.
is almost
to
In
straight.
This
behave
just
action
the
is
as
In
respiration.
higher
exerts
are
had
we
20
per
accelerated
grains
a
con-
toxic
much
in
expected
cent
5
m.
sugar
mol.
while in
firmer
will remain intact.
can
of
electrolytes
be ascertained in but
investigate
concentration
review, there
are
5
a
the action of
which shows
2,
a
cent,
ultimately, will
m.
n.
about the
examining the bursting of the grains (top
mol.
influence
Be(N0 3)
established in
per
amyl-alcohol
find them
burst.
10
20
respiration
expetiments regarding
the
observe
straight line but, here, alcohol
a
mol.).
m.
Fig.
see
shall
we
little while. This,
a
out.
In
we
interpretation
an
the influence of
measure
68),
Fig.
centration
allow of
to
37). Grains in
(Fig.
2
If
(Fig.
phenomenon
same
small
too
problem is aggravated by the fact that the grains
in many media.
on
as
is
measure
Our
as
that
high
a
as
must not
on
few
the
single
numerous
very
abt.
germination
salts.
Even
acid reaction,
.04
n.
of
cases, in bakers’
may
the
be
Out of
30 cations
be used since
they
are
toxic
62
in low concentration
(Ag, Cu, Hg, UO
and
inside
this influence is
the membrane;
internal system.
being
action
an
in
.6
living
and
Th(N0 )
internal system.
any
juice and that
do
To
that
again,
be put
the
on
some
not
and
we
down
living
in
part
in
change
a
to
to
make
Th(N0 )4
when
3
that where
to
this
AgN0
give
CuCl,
we
should exclude from
internal
§
2.
1.
2.
The
some
an
one
yeast
living cells though they
interpretation
this
being
membrane
protoplasmic
pollen there
are
further indication
The influence of
The
on
set
reason
our
3
)2
findings.
concentration
same
cells
a
of
induces
U02(N0
,
in.
Here,
too,
the
more
why
the five metals
acting
upon
systems.
In Lathyrus
give
on
organic
an
its action
CuCl,, HgCl 2
3,
experiments
our
action
an
connexion it should be
in the volume of the
in low concentration,
acts
yeast
changes
to
us,
the volume of the yeast cells
upon
trying
changes
by
of the
upon
the membrane,
to
Respiration of yeast cells is influenced in the
as
action
(78).
preparations
of the salts
use
In
upon the
act
acetone
permeability
this
be faced with
may
yeast.
antiseptics
influence yeast
not
does
58)
the action upon
decide whether the influence of
The influence of salts
us
naturally,
active
examine this substance both with respect
observed
(Fig.
demonstrate that the
attribute the results, established
we must not
must
30
it would appear
NaCl
4
concentration as
Since,
yeast.
take
cannot
substance should
some
the fermentation of yeast juice and
on
in the membrane. Once
an
eq.
much influence upon the volume of
exert
becomes felt in almost similar
juice,
an
upon
CuCl, after
discussion.
to
fermentation of
change
invade the yeast
to
From these considerations
Experiments by Warburg and Wiesel (98)
membrane
be
can
a
exerted
m.
time
some
3
influence of alcohols
yeast
as
to
decrease after
suddenly
needs
between
antagonism
lend itself
not
salts also
(Fig. 65).
the yeast cells
that the
the salt
Apparently,
cell. These five
by
caused
first, will
at
far
due, in the first place,
Respiration of yeast cells
fairly great
minutes.
is
it
not
As
Th).
a
ascertained,
as
two
to
of
Lathyrus
groups of
the
organic compounds
that may
of the membrane:
structure
on
pollen.
experiments
germination of the grains.
stability of the grains in different media.
To what
extent
do
these
bear
experiments
out
the
membrane
theory?
To
answer
of the
may
we
inside
this
question
we
phenomena occurring
assume
the cell
the
and
shall have
to
form
some sort
of
opinion
during germination. In the first place
existence
outside it,
of
a
variation in
essential
to
osmotic
germination.
pressure
When
the
63
osmotic pressure outside the cell is low,
When it
rapidly.
In
between
these
cent,
per
We
in each
the
two
there
is
region
a
takes
grains
pollen grains will burst
In
place.
germinate
not
of
at
concentration
this
experiments
our
all.
where
is
concentration.
sugar
If this
were
substance the
the
representing
the
grains will
imagine this mechanism
cannot
germination.
1.
high,
too
of the
germination
a 20
is
be the
to
only
controlling
one
then:
so,
bursting
germination and those
representing
curves
of
phenomenon
the
would
grains
coincide.
the
2.
curves
of all substances
coefficient)
It is
evident
consideration.
(allowance being made for the isotonic
would coincide in
that this
The
does
principle.
occur
not
germination
in
the
and
curves
bursting of the grains coincide, practically, in but
In
most
depression of germination
electrolytes
tration lower than that in which the
holds
representing
few
a
single
in
occurs
the
cases.
concen-
grains remain intact. The reverse
good for alcohols; here, grains germinate in spite of their being
When substances
intact.
may
under
experiments
those
be
expected
lie
to
toxic,
are
naturally, the germination
in front of the
curve
governing
curve
the intact
grains. The
be
cannot
The
in the
reverse
phenomenon, occurring in organic substances,
explained from the standpoint of osmotic pressure alone.
curves
of all substances
region of the
of the substance
would be found
of
cane
Hence,
on
lie
to
sugar,
cause
cannot
not
for
This leads
us
to
a
different
be
curve
a
toxic
which,
curve.
activity
naturally,
The behaviour
germination in higher concentration
explained
occurring
suffice
principle, namely
only
from this
hypothesis.
in osmotic
pressure
within
the cell
to
explain the effect of many compounds
to
be
seen
which mechanism is
actually
germination.
to our next
surmise which is
absolutely hypothetical
character:
A certain substance
within
the
cell
(possibly
towards
plasmic membrane. The
that
changes
when
in the cell
there exists
a
the
a
growth substance) permeates from
flanking cell
It, thus,
many
wall
distinct
(in
this
case
through the protopollen tube demands
the
intine)
occur.
difference between osmotic
becomes evident that
different
wall
formation of the
very
inside the cell and outside it, will the
i.
coincide in
And here, too,
in front of the sugar
germination. It’remains
responsible
ought
curve.
depressing
differences
and outside it do
in
might
methyl-alcohol,
than
sugar
grain be allowed
germination
may be
to
germinate.
depressed in
ways:
by substances acting
on
an
Only
pressure
internal system of the cell;
6
4
in
2.
is
essential
liquid;
germination,
to
checked.
The first
this
of the outside
permeation of the substance,
being
Hence,
strong concentration
excessivily
an
because
3.
methods of
two
shall be
we
allows
the
depression have already been reviewed.
concerned with the third method since
mostly
membrane
protoplasmic
to
assume
controlling
a
activity.
from
Firstly,
better
some
We may
freely
described
experiments
about this
of the
stability
take the
we
the
knowledge
assume
sugar concentrations
grains
that sugar
must
we
membrane.
in
different
hardly
that leave the
As
enters
alcohols
(Fig.
the cell.
Hence,
the osmotic
grains intact,
of these sugar solutions will
be the factor
wall of the
certain firmness, the
intact
pollen grain has
a
nation is
not
This makes it
depressed until occurring
in
in
value
grains will remain
yet, attained
as
possible
higher
a
34).
As the
deciding stability.
though osmotic pressure outside has not,
even
the value of the inside pressure.
obtain
to
try
starting-point
a
that
germi-
concen-
sugar
tration.
There
three
are
alcohols:
alcohol that stabilize
This would
the
mean
membrane,
the
that these alcohols
becoming
this
alcohol show
a
than there is
of sugar
with
This
action).
In
different
oleate
reminds
that the
faintly
or
action of
alcohols
In
the
A
some
an
butyl-alcohols
as
an
in
we
shall be
grows
entering
so
a
excessive
membrane,
This
faced
opening
coacervates.
must
a
slightly
This
be either
from the
sensitized
to
which
curves
(cf.
we
be
Fig.
36).
system
succeeded
is
a
shrinking
all
of
the
play
by
a
the
will
condensing action
an
opening
action.
Eventually,
we
grains
longer burst.
no
felt
explained along
butyl-alcohol
pressure outside the
in
is indicative of the fact
relations
be
coacervates.
where osmotic
electrolytes
may
Tert.
shall observe
that the latter will
symplex
an
exerts
pollen
This results
found in soap
region
The influence of
centration.
is
first
required
are
opening influence.
an
Lathyrus
strongly
example. At first,
minimum
we
upon
influence.
observe
us
Here,
phosphatide
at
have
sugar.
methyl-
and
symplex relations (i.e.
it will
a
opening
grains remain intact)
similar
the
membrane of
alcohols;
Ethyl-
More of these alcohols is
non-sensitized system.
exercise
firm.
propyl-alcohol
theory forwarded by
serve
(the
a
amyl-
n.
condensing action
a
of the
soap and
n.
shrinking action though later
suggests
and
butyl-
keep the grains intact.
us
coacervates
n.
exert
more
picture.
to
neutralizing action of
a
propyl-,
n.
grains in lower concentration than
comparatively
that,
in
principal
this
low
con-
protoplasmic
part.
(It
should
65
be observed
electric
reverse
part predominates
effect
to
that in the
change.)
a
Also
the great
We
conclude that this
finally
in
more
common
the
direction
of
for
points in the
pH
ioo)
grains
a
are
Jong
de
representative
of
that is
found in the
KC1
and
less
CaCl
Though
the
be
regarded
on
the
this
in
i.e.p.,
as
hypothesis,
the
too,
of
pure
7. The strong
H
same
direction
a
with Lathyrus
that
theory
the
complex system,
(cf.
we
lecithin
Fig.
may
indeed shown
assume
well
by
an
as
exert
21).
every
some
Basing ourselves
a
be
membrane should
that, in low sugar
KC1 will
will
70.
hope having shed
of this membrane model.
as
lecithin
pollen do not, in
protoplasmic
we
pure
antagonism between
Fig.
experiments
our
potentialities
are
15,
charge with CaCl2 than
69.
(15 per cent.), CaCIj
We
14,
number of
uni-complex of lecithin (Bonner)
the reversal of
The
pure.
points
2
evince
at
neighbourhood of p
Fig.
way,
a
(Winkler).
protein
lower concentration
lecithin
developed
(Bonner,
mainly of lecithin. This is confirmed by the fact that
a
have
Jong and Bonner’s model
The pn region, where
that the membrane consists
(6
—7),
designates
germinate
tri-complex
can
de
The three membrane models
potential models, ranging from
with
membrane will
protoplasmic
Bungenberg
Saubert, 30, 31, Winkler,
has
the
electrolyte will be needed
sensitivity
with Bungenberg
than with that of Winkler.
under
a
membrane in which
a
—
direction.
same
to
case
much of the
—
light
upon
concentration
stabilizing influence.
experimental series that KC1 and CaCl2
,
given concentration, increase the percentual germination in 15 %
will exert a more powerful action
sugar (Fig. 69). Naturally, CaCl2
in
a
than
KC1.
Since
we
imagine
methyl-alcohol,
methyl-alcohol
we
an
osmotic action
expect
to
that, in low
will be needed
to
be the
sugar
principal
agent in
concentration,
depress germination.
much
This indeed
5
66
be
to
appears
Less
(Fig. 70).
so
methyl-alcohol is needed
germination in high sugar concentration than in
The
§ 3.
membrane of
protoplasmic
Contrary
the
to
As
problem.
the influence
of alcohols
changes
we
As
in
pollen,
our
phenomena
first
effort
in
occurring
important in the solution of
with
the
proteins
the exterior,
the
cell.
meation
of
opinion,
our
glucose
Ni(N0 )
3
bear
out
2
so
far
as
exposing
they
are
chiefly concerned
are
permeate towards
cannot
(glucose)
must
it
highly probable
seems
be
can
they
as
either
presumption
permeate into-
necessarily
depressed
that
this
stimulated
or
per-
by
an
difference between the influence
vast
respiration and that exerted
on
this
problem. We
our
action upon the membrane. The
of
in
the
approach
to
directed toward
fermentation,
moreover,
the substrate
In
have
to
with
experiments
permeation of a substance takes place,
actively engaged in the formation of hexose
are
(77), and,
phosphates
shall be
attributed
whether
question
As
not.
or
yeast.
be
not
may
discard any
must
organic substances. Consequently, we shall
problem from an entirely different angle.
the
sugar.
Lathyrus pollen, here the eximportant in solving the membrane
are
in the membrane,
baker’s
depress
to
cent,
per
with
experiments
periments with electrolytes
20
permeation
on
seems
to
(Fig. 66).
When
reviewing the fermentation curves occurring in salts that
depress fermentation to a certain percentage of the blank (Fig. 53,
54 and
55)
we are
It suggests
faced with
a
Once the yeast is saturated,
influence.
part that is
conspicuously
horizontal.
saturation of yeast with the salt in low concentration.
a
Ultimately,
a
further addition of salt
in osmotic
will
exert
concentration fermentation
no
will
decrease.
If the yeast be
existence of
concentration,
actually saturated with
the
at
(Naturally,
this
saturation.)
If the
concentration,
the
AgNO
suspensions.
In
a
ation
our
g
As
concentration
we
then
we
of
a
a
is
different
concentration of
graph
are
In
Fig.
we
use
indeed observe
a
will
solution.
the
point of
with respect
curve
when
to
altering
and 7 per
cent,
substance that is active
yeast
in low
be observed.
caused
4 per
salt
71 the influence of both
cent,
normal the actual
presumably
experiments
shall
curve
1
the
the
beyond
different
a
a
the
considerable
compared with 5 per
of fact, in
assume
taken in and
might
we
of
extend
not
bound be
shall observe
matter
salt,
quantity
equilibrium,
does
strength of the yeast suspension.
KC1 and
In
point
assumption
amount
a
between the
equilibrium
an
cent,
fresh KC1
by
an
curves
glucose.
curve,
in the ferment-
osmotic
If
we
shifted
phenomenon.
raise this
to
12,
conspicuously
67
to
the left.
effect
Strengthening the sugar concentration, however, has little
upon
the
AgN0 3
amount
of cation bound is
brium concentration. This is
differences,
resulting
from
a
only be watched in salts acting in
can
low concentration. With respect
the
The
curve.
varying yeast concentration,
to
salts
small
acting in high concentration
compared with the equiliclearly demonstrated by Fig. 72. Only
as
acting in low concentration may we observe a fresh curve
altering the yeast concentration; in the “toxic” metals Ag and
in salts
when
Cu
variations
greater than in
are
non
Fig.
71.
Fig.
72.
toxic
metals
From these observations it would appear that
cations
our
has been
replaced by
starting-point,
of, for example,
another.
the Ca therein
Ni(N0 3 )
2,
the Ca
can
Taking
be
one
a
(e.g. Ni).
of the membrane
Ca
replaced.
tri-complex
tri-complex will slowly pass
The latter is denser than the Ca
Ni
tri-complex (cf. Fig. 54).
plex; both permeability and fermentation decrease. As
a
has
been
completely
replaced
by
Ni,
as
On the addition
fermentation
soon
will
into
com-
as
Ca
remain
68
to
Our
zero.
KC1
(cf.
membrane.
Ca may be
K
may
At all
times
as
strong
number
to
be
Mazia arrives
It is very
able
is
as
the
Mazia’s
want
be
directly
only
rate
be bound.
substitute
to
Na
approximately
is
In
a
Mg this
required.
conclusions: Ca forms
compound
organic substances showing little mobility.
likely that Ca
regarded
we
must
Of Ba and Sr less
20.
following
of
of Ca will
that of the latter substance.
as
is enclosed within
this substance where it
to
This
groups
If
very weak
not
its
solutions;
amount
concentration
roughly
the
at
itself is
binding
equal
an
the Ca from
expel
notably from
more,
The
by other cations.
their
Ca,
hundred times
with the acid
the
increase.
cannot
2
replaced
ought
in
permeable than the
more
the concentration of CaCl
for
ions
water
once
solutions.
2
it.
add
we
the Ca
be effected with the aid of K citrate.
can
be taken in
CaCl
to
on
for
similar
a
with distilled
rinsing
proportional
or
with
us
This, however,
mol.)
depends
reduced
Mazia
example.
(75)
binding of Ca through the protoplasm of Elodea. Even
Afterwards, Ca
(.00005
substituted
fermentation will
Here,
presents
established the
protoplasm.
is
tri-complex will be
but the K
fortnight’s
K
Even then,
48).
Literature
a
be
picture will be quite different if, for example,
Fig.
membrane,
original
concentration fermentation will
In osmotic
stationary.
outer
be
can
a
a
membrane
imperme-
readily replaced by other ions.
layer of the protoplasm.
fit quite well into our train of
thought. What he calls “a compound of Ca with organic substances
Obviously,
experiments
having little mobility”,
in
our
opinion
should
be
referred
complex system. Especially the ready exchangeability
points
in
this
substantiated
Boas’
direction.
from
our
concurrence
with
subsequently,
things, this
2
less
salts
this
a
pressing
action,
will
in
will
to
(condensing)
mixture
a
is
agent.
Since
of A12 (S0 4)
3
more
be
(9).
likewise
are
action
on
the
permeable.
In
of the salt which,
system
active,
Ba
a
(amongst
membrane
from
the
acting independently,
saponin
and
a
further
reduction of the volume).
distinguished
The salt,
an
exerts
latter
penetration
a
be less
salt.
action.
the
internal
an
be observed from
reverse
a
obtain
upon
might
foregoing pages,
Saponin
makes
as
exercises
an
other
With
being
previous
is
a
de-
opening
saponin greater fermentation
occur.
It should
cation
to
act
we
phenomenon
permeable
by
that
NaCl
will
may
in the
to
standpoint.
membrane cholesterin
hypothesis
our
with radio-active substances
by experimenting
experiments, referred
explainable
BaCI
Perhaps
to
of the cations
be
possible
to
replace
depressing fermentation,
establish
the influence
of
the Ca
upon which it
a
cation
in
the
ought
exerting
a
membrane
be
by
a
equally easy
stimulating action.
to
69
of yeast
Seven grams
On
the
Now,
we
coincides
the influence of KC1
with
will
yeast”
The maximum
that
be
by
least in part,
at
With
like
tri-complex
this
KC1,
resulting
of non-treated
Mn. This
when
that
Mn
This
(Fig.
KC1 raises
yeast;
between
similar
to
high valency
a
low
has
a
antagonism
stronger
results will
than the
models
If
were
be less favourable,
as
Fig.
Ni(NO )2
.2
a
we
have
has
pointed
influence
an
been
complex
to
the great
powerful
this
influence than
An anion with
one
possessing
K SG
observed;
be
may
2
shows
4
forces in the electric
a
KC1.
our
membrane.
Here, contraty
(Bonner).
with
importance
opinion,
In
seems
the series
of
less
Lathyrus,
to
constituted
are
part of the
sen-
potential
§) the protoplasmic membrane of yeast
closely the tri-complex (Winkler) than the
that govern the situation
cells
occurs
pre-treated
antagonism of the ions.
erythrocyte
more
K
demonstrates that in
76
m.eq.,
yeast which
by
the
(cf. previous
proximates
a
peat
experiment concerning the antag-
The membrane of the yeast cell, in
sitized
ferment-
75).
an
and KC1.
than
on
observe
we
treated yeast.
non
2
too,
Mn,
way
more
Here,
valency.
in
for
(Fig.
in the
valency
a
K
blank
.
of the anion
depressing action
replaced,,
fermentation above
Ni(N0 )2 In Chapter I, § 3,
3
a
of this
explanation
same
concentration of
that shown
action of KC1
in the
Ni(N0 )
3
constant
The
3
cm
the normal
under
lie
to
too
(Fig. 73).
this yeast
on
n.
is twice
74). Ni yeast is affected
closely with
corresponds
onism
a
MnCL2 .6
in the membrane has been
exerts
occurring
2,
die in this substance
as
3
stimulating
yeast.
substituting
with CuCl
experiment
found
from the
phenomenon is obvious. The Ca
ation.
cm
too
Later the yeast is shaken with
centrifuged.
measure
The blank of “Mn
blank.
day in
a
morning of the day of observation the suspension
diluted and then
water.
left for
are
the chief
apuni-
factors
by complex relations (the
membrane).
We have
been able
not
investigate the symplex relations in this membrane. This is due
to
a
to
phenomenon of the process
particular
Here, the action of alcohols
much
we
sooner
wish
refer
to
the
investigates
alcohols.
until in
less
than
It
Evidently,
where
of the
to
action
the
appears
an
that
carried
of invertase
latter
do
In
this
respect
by Kisch (68). He
out
the addition of various
on
not
influence
this
process
high concentration and this, again, suggests
a
membrane.
our
model
offers
an
explanation
protoplasmic membrane. We wish
our
becomes evident
the membrane.
upon
the
under consideration.
internal system
experiment
exosmosis
exceedingly
sensitized
of the
an
on
membrane model
may
be
to
cite
of many
a
few
properties
more
examples
instrumental in the solution
permeability problem. Thus, the glomerulus membrane (54)
70
Fig.
Fig.
73-
74-
Fig.
Fig.
76.
75-
71
is
impermeable
glucose only in
to
definite Ca
a
the Ca concentration is either stronger
will pass
looking
be
a
this
at
Certain
plants
In
red
the
large
according
of
Magistris
organic
check
to
to
When
case.
membrane
to
far
as
go
dyes
as
the
destroys
treatment
phenomenon. On the other
in KC1 solution,
occur
(i n.) gives rise
2
placed in distilled
are
partly built up from lipoids.
check this
dye-stuffs will
the
to
some
to
in
even
exosmosis;
i
n.
this,
hypertonic haemolysis
(too).
(72), in amplifying Hansteen-Cranner’s experiments,
the factors that define the exosmosis of
from
phosphorus
stimulate this
plant
organic
cells.
Also
P while
liberation. He finds
though
of the membrane.
The
Ca
2
a
to
appears
less
to
extent,
the action of CaCl2
in KCI.
not
anorganic and
CaCl
here,
KC1 and,
2
conception
this
may
this
is
know,
irreversible in distilled water,
can
they
Evidently,
we
required
the exosmosis of
MgCl
in this
expect
this
example,
views, may be likened
to our
erythrocytes
gets down
as
of CaCl
amount
we
when
lipoids
for
cells.
membrane which,
hand exosmosis of
important
system.
secrete
But little CaCl2 is
A
standpoint,
our
beetroot,
from
permeating
cell
from
unstable
very
water.
will be very
pH, too,
through.
concentration. As
weaker, part of the glucose
or
to
This confirms
be
our
taken in
only be exchanged for another cation,
by the membrane
example K.
for
The practical
work was carried out in the laboratory of medical
chemistry of the Government University at Leyden. I am greatly
indebted
to
who
given
has
its Director, Professor
me
the
Dr.
ungrudging
H.
help
G.
Bungenberg
of his
Jong
de
and
expert advice
valuable criticisms.
SUMMARY.
As
as
to
the
teresting
the
forms
present
The
of
part
cells
of the
etc.
for
satisfactorily
a
layer
protoplasm
the
suggest
that
though
by itself,
the membrane
this
being
layer,
properties.
of protoplasmic
membrane developed
explain but part of the facts. They concur
permeability
compounds,
explained
in-
facts
which stands
different
doubt
would be
On the contrary,
permeability theories,
Since
many
not
will possess
can
Numerous
structure.
of the
models
numerous
one
theory,
its
no
it
membrane,
protoplasmic
protoplasm.
the surface,
(47, 60, 70,74, 81, 82)
with
a
membrane is
protoplasmic
quite independent
on
of
investigate
to
actually
of Chambers and Reznikoff leave
experiments
the existence
from
the sieve
of different
investigated
a
single
theory, the lipoid
plant
previously,
theoretic
and
animal
cannot
standpoint,
we
be
shall
72
have
look for
to
To
Bungenberg
this
to
the
distinguished.
by
the attraction
In
the other,
forces
Firstly,
in the membrane
medium,
shall
of life.
We
caused
by the addition
now
the influence exerted
(if
gives
we
us
some
discard the
in
system
We
bakers’
compounds
20
As
per
sensitive
is
this
of
amount
with
the
in
cell
particles
the
contacts
many
processes
process,
compounds, with
membrane models.
on
of the membrane
upon
internal
an
the
burst
Also
(Lathyrus
influence
and
pollen
of
different
processes.
is
pollen
in
depressed
distilled
shall
we
the
During
Lathyrus;
in
region,
two
our
pollen
where
we
a
When
be
to
exerts
pn
summers
the
we
to a
amount
of Ca
w
;
ll
readily
an
has
difference
summer
is borne
active
an
in
since
out
the
part
behaviour
by
on
We
of the
antagonism takes
a
may
anticipate
alkaline
earth
a
less
metals
in-
the
marked
in
either
less active part here. This, indeed,
the facts.
Organic substances
exerted
in this.
the
cause
important variations in the alkali salts since the antagonism of
ions
in-
have
varying
electrolyte in the membrane in either year. Even
in the relative
very
no
sequence of
attribute
of
constant
experiments.
find
year
This
different sub-
by
water,
successive
either
presented great variations.
appreciable difference
the
note
biological
a
nature
objects
particular
of salts,
(6—7).
however,
compound acting
a
the
membrane).
in
to
the
will have to be used in all
action.
limited
to
as
As
we cannot
between the
affect
substances
same
of their life
the influence
experimented
alkali salts
more
membrane.
we
the
may vary. The proto-
too,
changes
alter
may
molecules,
can,
where
different
two
grains choose
to
fluence,
the
to
Lathyrus
cent, sucrose
measuring
by the
some
Germination of
stances.
the
Waals
der
given object,
the medium of various
to
eventuality of
and
on
We
region
compare
examined
yeast)
the
the membrane will
information
preference
have
within
changes.
the
being
governed
part
(complex relations).
substances
When the interval
inside
change
a
membranes
London-Van
various
changes, permeability,
membrane
plasmic
of
answer
leading part.
relationship
these
measure
ready
of these
greater thickness than a few
no
of these changes.
outcome
This
that
direction
a
electric
an
the membrane in any
the fact
complex and symplex
membrane has
the
the
play
study
to
of
use
immediately
section
hydrophobe,
(symplex relations)
make
is
between the ionized groups
When wanting
can
different parts
there
the
100) present
15, 30, 31,
theories.
permeability
under
developed
entirely
Two
all of the
comprising
models
Jong (14,
de
question.
be
can
model
a
mind
our
lipophile
come
up
to
coacervates.
our
expectations of the influence
There will be
analogy between
a
73
series of
activity
organic compounds, placed
oleate
on
(with the exception
Pollen
influence
an
even
Also alcohols
of
will
grains
value of the
show
this
surroundings
cane
that
be observed in
may
conclusion
sensitized
If
in
all
at
it
relations
are
opinion,
the
Jong
be
1.
stabilize
not
this
From
it
the
would
grains
appear
opening action while propyl-,
exert an
A similar
This
coacervates.
clear
active
protoplasmic
and
membrane
a
developed.
new
also
phenomenon
leads
must
to
us
be either
salts that
membrane,
relations.
complex
(a uni-complex
the
non-
Lathyrus
developed
method of
pollen
by
in
our
conforms
Bungenberg
of
the
measuring
we
have
rate
of fermentation
experimented with,
may
three groups:
depress fermentation
completely
symplex
Hence,
lecithin).
glucose by bakers’ yeast in different
The many salts,
divided into
roughly
this
membrane of
the fermentation of
trace
in
that,
than
the membrane model
to
Bonner
surroundings,
was
grains
Comparing them with
weakly sensitized.
becomes
more
pre-eminently
To
sugar
cane
high osmotic
great sensitivity for electrolytes and variations
antagonism of the ions and the influence of organic
the strong
compounds,
de
but
or
do
reached.
the
consider its
we
pii,
oleate
exert
for the fact that
shrinking agents.
are
salts
too,
Since
assume
responsible
protoplasmic
that this
action.
may
ethyl-alcohol
methyl- and ethyl-alcohol
butyl- and amyl-alcohol
diminishing
germination
(the grains remaining intact).
concentration.
concentration is
higher
we
Here,
media.
stabilizing
be
to
sugar
and
methyl-
sugar,
until
high
depressing
variations).
in low concentration
will
in
series
in certain
hardly permeates into the cells,
remain intact
the order of their
n
a
minor
some
burst
i
and
coacervates,
in low
concentration
(Ag, Cu, Hg, U0 2 and Th);
2.
initially lower fermentation
salts that
blank.
tration
They do
is
not
increased.
exert
to
a
given percentage of the
further influence
Ultimately,
abt.
at
i
when the
n.,
these
concen-
salts
depress fermentation completely (Tl, Be, Zn, Cd, Pb, Mn,
Ni,
3.
salts
Co,
Al, Fe
-
•
•,
La and
diminish
The first
(These
Ce);
that stimulate fermentation,
in low concentration. Later,
will
Fe' *,
those that
or
about
exert no
influence
fermentation will
n.,
again
(Li, Na, K, Rb, Cs, NH4 Mg, Ca, Sr, Ba, luteo and hexol).
at
1
,
group
salts
of salts, it
appears,
acts
have great influence also
upon
on
the
an
internal system.
volume of the
yeast
cells.) The influence exerted by the second and third groups of salts
can
be
traced
membrane
to
back
be
a
to
Ca
one
and the
tri-complex.
replaced by another cation. In the
same
principle.
Now, if
one
case
we
add
the
We
a
assume
our
salt, Ca will be
tri-complex formed
74
(for example
membrane.
Ni
a
In
tri-complex) will be less permeable than the original
other
which
to
In
been added.
the other
faced with
Upon
retarded in
re-
K
a
tri-
the membrane gets
concentration of abt.
a
volume of the cells;
we
are
phenomenon.
salts
respiration
is
process
osmotic
an
(e.g.
case
be stimulated since
parallel with the diminishing
runs
fermented
glucose
might be fermented by yeast
permeable. Decreased fermentation in
more
n.
has
of
amount
that
amount
fermentation will
complex)
i
salt
no
the
words,
presents but part of the
have
quite
a
influence.
different
concentration which is
a
This
practically identical
that in which volume decreases.
to
Alcohols
discard
wanting
In
presumably
the
arrive
to
this
results
upon
conclusions
at
an
with
and Ca
these
hence
we
must
when
compounds
regarding the membrane.
membrane
This model will be
lecithin, protein
internal system,
experiments
protoplasmic
dominant part.
are
act
of
relations
complex
a
play
a
pre-
tri-complex whose components
(Winkler’s
model), possibly accompanied
by substances which intensify the symplex relations (“sensitizers”).
We
look
must
Bungenberg
extensive
series
upon
of
the
and his
Jong
de
membrane
collaborators
This
potentialities.
models
as
series
mere
ranges from
to
(Winkler).
these
component of
systems
Every
the
construed
from
one
few
a
a
of the
structure
in
These
data.
single
which the membrane is
models
with
reconcile,
show
It
expected
various
be
cannot
would become necessary
those
on
to
biological
be active.
to
distinct
following
the
another,
one
lecithin
be varied.
may
membrane
protoplasmic
investigate the influence of numerous compounds
processes
a
tri-complex of lecithin, protein and Ca
uni-complex (Bonner)
Evidently,
developed
by
examples of an
advantages:
permeability
they
theories,
and,
alike, they will allow the differences in the permeability of organisms
and organs
ponents.
to
Our
be attributed
great lack
of
to
variations of
knowledge
properties of the membrane components
we
can
no
existing
among
problems,
in the
The
for
than
at
protoplasmic
example
least for
the
accounts
most
remarkable
sugars
the present,
for
the fact
important
membranes of
the very
com-
physico-chemical
various
that
differences
cells.
differences
Minor
occurring
through glomerulus membrane,
be solved.
general importance of complex systems, possessing, amongst
lipophile components, is clearly shown by the cir-
things,
cumstance
that
two
the series
of
different objects like
Lathyrus pollen
protoplasmic membranes that can be included
vastly
and bakers’ yeast have
in
construe
permeation of isomeric
cannot,
other
more
the membrane
about the
potential
models without great
difficulty.
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1.
Adam,
N.
2.
Baas
The
K„
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and
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Chemistry of Surfaces. Oxford Univ.
1930.
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L.
G.
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