Ada
Botanica
The
8
JVeerlandica
Influence
on
of
Growth
of
1-39
(1958)
the
Red
and
and
Avena
G.
Far
Red
Light
Phototropism
Seedling
Blaauw-Jansen
(Botanical Laboratory,
Utrecht)
December 5th,
(received
1958)
CONTENTS
chapter
Influence
chapter
it.
chapter
in.
of
Determination
in
connection
the
vi.
Extraction
chapter
vii.
Identity
chapter
viii.
The
The
of
its
to
light
of A
....
=
660
m/i
11
metabolism
substance
growth
10
length
wave
coleoptile
...
of the I.A.A. content of irradiated
and
16
coleoptiles
the
the
R.L.F.
Avena
ix.
of
effective
most
Avena
with
non-irradiated
chapter
chapter
the
of
4
light
Quantitative analysis
chapter v.
1
methods
orange
The reaction of
iv.
chapter
and
Introduction
i.
in
“Red
Red
Factor”
Light
18
(R.L.F.)
Factor
Light
24
connection with the
of
reactions
phototropic
27
coleoptiles
far
influence of
red
(of A 740 m/i)
light
on
the Avena
32
Coleoptile
/“N
1*1.1
1*
P
Consequences which the discovery
chapter x.
red
by
the
and
light
orange
of
application
on
auxin
of
Avena
. 1
•
/I
*1
the influence exercised
will
coleoptiles
have
for
36
tests
summary
39
references
39
INTRODUCTORY
Red
light
activates
or
the
perceived by
an
Red
(R.L.F.).
the
is
also
reduces
coleoptile,
and
increases
At
low
I.A.A.
blue
the
light.
curve
primary
which
unidentified substance,
light
the
the
first
as
positive phototrophic
the
function
a
seedling,
termed
we
the
of indole-3-acetic
content
concentrations
presenting growth
REMARKS
leaf of the Avena
R.L.F.
of
the
into
R.L.F.
light
and
I.A.A.
growth-substance
of
the
light
coleoptile,
on
the
The
tests
but
were
it
formation
of the
implications
studied.
counteracts
of
the
Far
red
and
use
of
light
too
the
growth
reduces
interaction
for
in
induced by
two-peaked
a
the
factor
(I.A.A.)
rate:
which
curve.
of the
the commercial
the
I.A.A.
content
eventually annihilates the effect of red
R.L.F.
CHAPTER
I
INTRODUCTION AND
1.
red
to
produces
concentra.ion,
normally shows a single peak, is in this way transformed
“disposition” phenomena in phototropism are due
The
acid
curvature
enhances
I.A.A.
and
red-light
METHODS
Introduction
The dark
made
rooms
usually
are
in which
lighted
experiments
with
orange
1
with Avena
or
red
light,
coleoptiles
although
are
it
is
2
G.
known that
the
According
of
[effect
Vogt
early
as
Bonner
some
these
to
way
It seemed
growth,
at
first
but
not
phototropic
at the
regions of
tropic
to
To
that
due
to be due
this
for
end
to
This
but
possibility
this
influences
the
in
a
phototropic
in
spectrum,
formed
of
in
phenomenon
have
not
obtained.
induce
not
curvature,
a
blue
to
response
series
a
this
photo-
rough estimate of the
a
actually
was
a
growth
a
showed that the latter could
that
rate
as
For
was
actually
were
light, but
of
rate
coleoptiles.
coleoptiles
curvatures
the size
investigated
was
by
and
coleoptiles,
On closer examination this
too.
the
Liverman
whether there
placed
we
in
illuminated
suggested that the red light itself did
that it
found
Konings-
by
light would affect the
difference
a
scattered blue
of
curvature
a
(1941),
response of the
ourselves
intensity of the scattered light
caused
that red
to
phototropic
the red part of the spectrum
appeared
Schneider
unilaterally
see
confirmed
was
this effect.
was
(1957).
phototropic
sides of
response.
observed
effect
sight unlikely
decided
we
this
light
investigators,
confirm
not
red
(1920)
later
growth of coleoptiles
Lint
is
could
light,
(1937),
the
but
the
on
De
curvature
opposite
reason
and
Avery
Zollikofer
to
coleoptiles,
red
purer
light
and
(1953)
the
on
1915, and
as
(1922),
berger
a
red
and
(1909)
effect
used
probably
An
Blaauw
to
phototropic
a
who
2.
in
respond
spectrum.
has
of
do
coleoptiles
BLAAUW—JANSEN
This
light.
described
experiments
in
paper.
Material
The
and
general
experiments
“Siegeshafer”.
them in
in
a
at
a
in
moist
dish;
petri
temperature
kept
at
of 75
%.
and
water,
tap
closed
grains
same
or
in
plants
hours
holders
glass
88
were
under
the
in
of 22° C
hours
at
old,
an
grains
dark
a
and
sativa
by
moistened
on
placed
24
Avena
wetted
thoroughly
was
After
of
seedlings
germinate
to
put
temperature
When the
with
were
the latter
of 22° G.
vermiculitc
the
made
were
Unhusked
methods
a
they
filterpapcr
orange
lamp
planted
were
which
room
relative
were
c.v.
shaking
was
humidity
used for the
experiments.
solutions of indole-3-acetic acid
All
made
with
acetic
tap
light
Watt) by the
produced by
with
a
set
an
0.5
by the
obtained from incandescent
incandescent
an
of
cell
a
(60
parallel beam and with
Tungsram.
of
in
Watt)
Moll
(Kipp,
sensitivity
was
an
inter-
of which the tolerance
higher intensity
light intensity
The
light
in combination
Watt)
parallel beam from
3000
Volt,
thermopile after
from
a
light
similar interference filter. The
means
get
approximatively
lamp
100
lamp (12 Volt,
to
lamps (220 Volt,
Monochromatic
by Balzer (Liechtenstein),
Monochromatic
of
incandescent
layer
(I.A.A. or heteroauxin) were
mg/1 KG1 and 0.2 ml/1 glacial
of Schott OG 2 filters.
of lenses in order
%.
use
was
use
ference filter made
was
750
containing
acid.
The orange
20
water
a
was
obtained
more
powerful
combination
was
Delft)
of the
with
measured either
or
with
a
thermopile
a
by
barrierwas
not
THE
INFLUENCE
sufficient
fore
for
these
the
Delft)
no
an
of the
than
exact
is
length
wave
tive
such
example of
± 8 m/t.
and
of
table
the
on
1.
Fig.
A
of the
intensities
light
page 4.
Example
The
of
a
light
that
transmission
the
mean
error
of the
length by
Zernike
Therebarrier-
of the
means
of the
galvanometer
large variability of the plant
%
was
aimed
are
meant;
and
1
Fig.
half-peak
sources
is
obtained
a
spectra
representa-
width in each
filters
obtained
spectrum
at.
transmission
mentioned,
were
values of the curvatures,
number of
coleoptiles.
a
intensities.
means
given in the tables
figures
ference
in
a
3
AVENA SEEDLING
light
wave
The
spectrum.
a
summary
10
THE
by
out
In view of the
precision
ON
lower
instrument
obtained with the interference filters
is
LIGHT
calibrated for each
used.
was
RED
carried
were
indicator
an
greater
Whenever
FAR
measurement
was
As
thermopile.
(Kipp,
AND
measurements
layer cell which
material
RED
OF
are
on
by
that
were
case
used
in
given
the
experiments
means
of
an
are
inter-
filter.
or
of the
The variation is
rate
of
growth, measured
expressed
as
the standard
mean:
SE
~
m
'
n
=
]£(»
the number
—
m
)
2
=
the
of
coleoptiles
sum
of
Sometimes the results
the
are
n
[n
(always
squared
—
1)
mentioned between
deviations
given in the form of
a
from
brackets)
the
mean.
graph. The figures
4
G.
Light
Light
BLAAUW-JANSEN
source:
Filter
incandescent
bulb
Light intensity
in
colour,
wave
resp.
tension
energy
length
in Watts
in
Volts
20
220
Schott
75
220
Schott
sec.
>
550
A
at
400
m/x
520-620
400
m/x
m/x
480
560
100
12
3000
60
int.
f.
477
m
100
12
60
20
f.
int.
562
20
100
12
int.
f.
586
600
100
12
int.
f.
595
620
100
12
int.
f.
614
640
100
12
int.
f.
644
660
100
12
int.
f.
657
m/x
m
int.
f.
int.
f.
682
m/x
700
100
12
int.
f.
702
m/x
740
100
12
int.
f.
741
12
int. f.
drawn,
1.9
m/x
12
were
1.4
mju
60
graphs
0.9
/i
100
the
0.5
m/t
680
from which
0.6
m/(
int. f.
580
3000
0.5
fi
int. f.
3000
3000
a
maximum
Orange
in
2
kind
Green
A
ergs/cm
transmission
2.1
70
2.9
2.3
2.5
m/x
50
are,
always
moreover,
given in
table.
The
following general policy
derived from the results
was
kind,
did
we
ments,
but
Where
by
this
to
try
to
method
of
its
two
presumably
of such
a
Experiments
the
the first
size
blue
of
on
place
the
of the
same
same
experi-
be
series
difference
are
to
above
preferred
by greatly
be
an
increasing
experiments.
INFLUENCE OF
In
conclusion
a
experiments
experimental
should
CHAPTER
1.
whenever
validity by repeating the
between
error
three
or
implications.
differences
reduce the
the number
its
test
checking
small
expected,
attempt
not
pursued:
was
of
II
ORANGE
LIGHT
phototropism
we
investigated
curvature
whether orange
induced
by
unilateral
light influences
illumination
with
light.
Method
Germinated seeds
were
downwards, in vermiculite,
darkness, except
been
When
the
transferred,
The
in
placed
for
the
plants
placed vertically,
8 in
period of
ten
vermiculite,
at
a
were
88
also in absolute
coleoptiles
were
a row.
hours
The
with
the
embryo pointing
were
kept in absolute
minutes 46 hours after
they had
which
watered.
old,
darkness,
exposed
with
plants
to
their
time
each
they
series
were
of
8
the illumination
narrow
side
plants
was
apparatus.
to an
unilateral
THE
INFLUENCE
illumination
at
once
with
AND
different
after
or
RED
FAR
LIGHT
of
amounts
THE
ON
with
480 m/r,
=
of
light
5
SEEDLING
AVENA
of A
light
illumination
previous
a
RED
either
different
a
length.
wave
1\
after
hours
coleoptiles
and
OF
were
illumination
the
severed from the
The
shadowgraphed.
with
curvatures
of
light
seeds, placed
A
measured
were
480
=
by
the
m fi
photographic
on
paper,
of
means
a
protractor.
Experiments
whether
Coleoptiles,
ergs/cm
2
orange
all
In
induced
Table I the
The
results
of
first
a
larger
positive
I
of
amounts
with
light of
36
X
have
been
TABLE
A
=
A
=
in
480
Not
m/x
ergs/cm
two
experiments
in
plotted
given.
are
2.
Fig.
I
in
4
36.10
pre-illuminated
curvature
2
degrees
(6)
±
1.6
(8)
3
12.8 ±
1.1
5
13.0
1.2
1
1
0.5
6.3 ±
2
10.4
20
12.0 ±
120
II
5
10
20
30
From
these
the
orange
in
that the maximum
really
seemed
Now
the
a.
might
The
tion
In
that
be
of
explained
coleoptile
with
very
under this
therefore
b.
relatively
case
pigment
with
9.0
±
0.9
(7)
(6)
16.4
±
1.4
(8)
1.0
(7)
± 0.3
(12)
22.9 ±
1.0
(8)
8.0 ± 2.5
(4)
7.7
1.0
(6)
1.0
(9)
21.2
±1.0
(6)
14.0
±
±
0.9
(7)
23.2
± 3.2
(5)
13.7 ±
1.0
(7)
23.5
± 3.5
(4)
12.5
1.4
(8)
19.4
±1.8
(5)
±
appears
that orange
of the
enhanced
light
from
14
such
a
degrees,
23
to
undoubtedly
in
coleoptiles
way
which
why
the
by
small
be
curvatures
the
aid
with
of various
a
in
very
the
and
amounts
could
of
no
be
orange
small
observed
are
red
or
orange
after
light.
suppositions.
amount
coleoptile
is
the shade side
of orange
expected
light.
curvatures
are
It
considerable,
already reached
not
is
light [are administered.
only
after
appeared
illumination
however
obtained. This
that
supposition
discarded.
light
as
too
no
coleoptiles
of the pigment
light. Consequently with unilateral illumina-
curvatures
condition
Orange
(9)
(8)
±
contains but
low intensities
may
0.8
21.1
the
light-intensity gradient
light saturation at the light
when but
±
±
arises
that absorbs the orange
and
(8)
9.0
surprising.
question
unilateral illumination
This
4.7 ±
(10)
curvature
value is
degrees
0.8
14.0
it
experiments
phototropic
2
light
1.1
8.5
1
influences
±
with
ergs/cm
curvature
1
4
480
Pre-illuminated
Light quantity
No.
Exp.
10
illuminated
were
curvature.
measured in
curvatures
experiment
above
pre-illuminated,
not
or
unilaterally with progressively
which
from
pre-illuminated
light
a
induces
the
consequence
formation
a
of
steepening
more
of
the
blue-absorbing
light-intensity
6
G.
in
gradient
2.
Fig.
may
be
irradiation
due
phototropic
The
begins
to
react
curvature
blue-absorbing pigment
of blue
after
the
case
illumination
recorded of
are
either before
there
is
orange
growth
of coleoptiles:
not
•
illumination.
coleoptiles
in
blue
The presence
that
Table
20 ergs/cm
20
ergs/cm
20
ergs/cm
A
2
A
A
orange
=
size
of
followed by
m/i,
from
it
the
by
could
be
shown
36
4
10
X
36
the
104
x
that
future
TABLE
Unilateral illumination with 20
ergs/cm
2
ergs/cm
2
orange
coleoptile
light
side
10 4
9.6
x
10
4
ergs/cm
ergs/cm
2
orange
2
orange
light
light
degrees
(10)
27.9 ±
1-1
(14)
28.5 ±
1.4
(13)
with
has
the
orange
same
effect
III).
III
ergs/cm
2
A
=
480
m/i
after
curvature
10.7 ±
x
in
14.7 ±
(Table
.
9.6
light
1.3
pre-illumination
unilateral pre-illumination
none
applied
curvatures
Curvature
shade side of the
from
was
the
curvatures.
above
the future
pre-illumination
preceded
more
amount
II
m/i
m/i,
of
larger
a
II
from above
480
light
Moreover,
light from
as
480
=
light
orange
2
480
pre-
light. It appears that
Unilateral illumination
—
O
pre-irradiated,
irradiated with
were
TABLE
2
photo-
quantities
the shade side
at
influence if it
no
light. In
which
the
positive
light
produce this reaction of the shade
to
light should have
with
afterwards
light.
orange
after the illumination with blue
or
difference
no
increasing
therefore would entail
light would be needed
side. In this
with
with
the
upon
illuminated
are
the fact that the
to
irradiated
too
they
The reduction of the first
light.
after
curvature
admittedly
when
coleoptiles
the
with blue
unilaterally
tropic
BLAAUW—JANSEN
from the future
from the future
in
0.9
degrees
(9)
light side
22.7
± 2.0
(7)
shade side
22.7
±
0.8
(9)
THE
For
INFLUENCE
these
induce
is
third way
the results
could be
sensitive
2.
we
AND
a
FAR
RED
consider it
the
number of
ON
effect
THE
that
unlikely
7
AVENA SEEDLING
light would
orange
pigment.
of orange
experiments
light
the
on
derived
was
place where the
perceived.
particularly
demonstrated that
to
LIGHT
blue-absorbing
a
explain
to
of
is
light
orange
It
reasons
RED
the formation of
A
c.
from
OF
of
tip of the seedling
light.
orange
Location
the
sensitivity
the
to
orange
light
Method
In absolute darkness
tin
foil
1914
of 4
caps
tips of the coleoptiles
the
the very
length,
supplied with
were
that Arisz
same
the influence of illumination
study
to
mm
had used in
the base
on
of the
part
coleoptilc.
Then
the
whereupon
with
supplied
were
order
obtain
to
a
dose
a
removed in darkness.
were
X
unilaterally illuminated with light of
were
in
coleoptiles
the caps
positive
first
480
—
curvature
of orange
Next
as
light,
coleoptiles
the
2
m
(20 ergs/cm )
fx
large
possible.
as
Experiments
TABLE
Unilateral
Exp.
illumination
Nr.
with
10 4
ergs/cm
9.6
x
10*
ergs/cm
28.8
x
10
5
ergs/cm
=
480
104
x
4
10
x
From Table IV
clear
that
2
Curvature
orange
light, without
orange
light, with
caps
orange
light, with
caps
orange
light,
without
orange
light,
with
2
2
ergs/cm 2
ergs/cm
2
in which the
orange
light
has
no
influence if the upper most 4
Since the
possibilities
another
but
the
It
conclusion
of
tip
was
22.7
caps
the
a
and b
than
resulting
of the
had
it
that
caps
curvatures
influence
mm
all
at
remainder of
Therefore
we
because
curvatures
The
in
it
passed
on
could
be
the
effect
or
seedling
degrees
to
be
is
not
rejected,
the
tip
to
the
to
of
orange
(7)
(7)
15.5
± 2.7
(6)
13.0
±
1.7
(7)
19.0
±
1.4
(12)
11.0
±
1.4
(H)
recorded, it is
but
a
very
slight
screened from it.
arrive
cannot
one
of the
the
coleoptile,
orange
confirm this conclusion
light.
by removing
the reactions
to
orange
plant.
the
application
shown
standard
to
(9)
1.9
are
are
primary leaf that responds
technically impossible
of the
in
± 2.0
9.7 ±
caps
primary leaf in darkness and by studying
light
after
m/x
none
24.6
3.
A
10.7 ± 0.9
X
24.6
test,
2
none
9.6
1I
the
ergs/cm
Pre-illumination
I
at
IV
20
Avena
that
test
light
on
cultivated
in
of the standard Avena
orange
as
influences
light
the
well.
the
standard
Avena
test
Method
Avena
chapter
I.
seedlings
They
were
were
illuminated with
glass
orange
holders
light
as
while
described
in
germinating
8
G.
filter paper and also
on
BLAAUW—JANSEN
in absolute
growing
the
relative
96
to
darkness, except
carried out in orange
was
test
light. When the plants
room
a
88
were
they
were
selection
hours
old,
raised from 75
was
%
%.
with
decapitation
in
tated
The
With
in
of
part
the
till
darkness
first
the
plants
the remainder
darkness;
in
light (A
decapi-
were
decapitation
the
solution
a
=
560
for
second time after
a
sojourn
an
m/r). The primary leaf was loosened
because this
manipulation could
be
not
darkness.
hours after
in
decapitated
were
green
after the second
performed
After
stumps.
remained in absolute
plants
orange light.
coleoptiles
soaked
12
decapitation.
performed
was
of 2 hours in
Two
their
of
moment
very
second
of
another
decapitation
I.A.A.,
hours
two
blocks
agar
placed
were
the
plants
3
mm
of 3.6
unilaterally
of the
shadowgraphs
taken. Between these actions
were
till the
on
the 72nd hour when
at
humidity of the conditioned
The racks
the
had been
during the first 24 hours after they
put in the glass holders. From the 48th hour
the
upon
plants
test
kept in the dark.
were
Experiments
curvature of
plants which had been decapitated in orange light
compared with the curvature of plants which had been gropingly
The
was
in darkness. After
decapitated
agar blocks
by
in
performed
were
the first
light.'In
green
various
containing
amounts
TABLE
I.A.A. concentration
in
•
0.50
•
0.75
•
1.00
1.25
2.00
It
•
■
•
io- 7
I.A.A.
9.0
10~ 7
io-
7
io- 7
In
was
degrees
in
Decapitation
0.5
(11)
±
0.9
(9)
(22)
11.4 ±
0.9
(19)
17.6
1.0
(12)
15.2
±
21.2 ± 2.4
(9)
13.5
±
0.5
(11)
±
±
7.8
orange
1.0
14.1
±
0.7
(23)
14.7
±
0.5
(22)
14.9
±
1.5
(10)
12.1 ±
1.2
(12)
that orange
10~7
light applied
of the maximum
from
g/ml)
perception
the
purpose
in orange
plants
the
plants
loosened
of
first
about
21°
but
seemed
it
the
the first
during
(at
curvature
about
to
a
was
of
effect
of
14°.
by
decapitation
concentration
suitable
light
orange
decapitation
a
light
(9)
7
supposed
case
in
dark
IO"
To that
of the
presented.
12.5 ± 0.5
is unexplained,
orange light
the
are
io-
the size
of
produced
curvatures
7
appears
reduces
the
Decapitation in
the
of I.A.A.
V
Curvature
g/ml
0.25
decapitation all manipulations
Table V
This
means
the
carried
to
check
primary
out
leaf.
with part
light and with the control plants in darkness.
decapitated in orange light, the primary leaf
immediately after the first decapitation or after
were
either
the second.
In
or
the first
case
the
primary leaf
three minutes after it had been
second
were
case
carried
it remained intact
out
The solution
in
green
of 10 -7
for
light
left
was
exposed
two
of
g/ml I.A.A.
/
intact
the
only during
orange
hours. Further
=
was
to
two
light, in the
manipulations
560 m^.
the
only
one
that
was
used in
THE
these
INFLUENCE
proved
It
the
be
to
It
is
left
light
intact
the
decapitation
the
in
in
in
dark
orange
light
after
experiments
of
the
the
all
of the
tip
all
solution
of
Curvature
degrees
12.9 ±
0.4
(24)
(32)
influence
exercised
showed
that it
responds
by
the
to
light
orange
should
be
on
particularly
light.This
orange
7
loosened.
carried
g/ml I.A.A.
used
was
is left
taken away
as
a
test
the
in
decapitation
X
of
560
—
m/(.
solution.
light reduces
intact
but
decapitation
the first
light
green
in
or
whereas
whole,
a
second
Except
in
out
light
orange
as
was
At the
left intact.
was
in
removed
was
tip of the coleoptile
was
were
1CV
either
performed
during
the
some
curvature
time after
irradiation.
decapitation
First
VII
Tip of the primary
leaf
Curvature
in degrees
in
the
dark
intact
16.1
±0.7
(35)
in
the
dark
removed
17.1
±0.7
(33)
in
in
4.
orange
light
intact
12.8
±
0.6
(35)
orange
light
removed
15.9
± 0.6
(33)
Conclusion
From
light
the
play
of
publication
elongation
the
tip of the
to
discussion
series
a
first
of
phototropic
via
the
special
Sharman
well
as
tip
We
a
and from
leaf
leaf
the
coleoptiles
leaf may
the
and
preceding
influences
of Avena
experiments
reactions
did
(1942),
come
who
a
is
to
our
observed
differentiation
of Roberts
leaf has
it
and the
clear
that orange
reaction
tip of the primary leaf. That
part,
the
as
paper
of the
in
tissues
(1951), who stated
larger chlorophyll
the
tip of
from
knowledge
that
I.A.A.
on
£ea
start
the
at
that in wheat
than the
content
a
leaves
rest
leaf.
did
not
orange
coleoptile.
coleoptile,
will
ourselves
occupy
light, but
consequences of such
the
in
(33)
that
the
pri-
light.
removed
TABLE
of
the
VI
From Table VII it is obvious that orange
a
is
0.6
only if the tip of the primary leaf
the
it
indeed
after
16.6 ±
tip
leaf
primary
manipulations
A
that
only
curvature
time
some
decapitation
2nd decapitation
seedling
the
primary leaf
the
series
the
reduces
follows.
the control series
tip of
of orange
0.7
decapitation,
the
effect
15.7 ±
leaf
primary
as
first
darkness,
in
the
9
SEEDLING
decapitation
the
on
with screened
checked
At
1st
after
tip
the
2nd
light
coleoptiles
was
after
orange
The
leaf
Primary
AVENA
the
during
orange
TABLE
First
THE
ON
concentration.
concluded
was
perceives
LIGHT
experiments
this
at
that orange
therefore
leaf which
mary
RED
FAR
preliminary
leaf
primary
irradiation.
AND
largest
(Table VI)
appears
when
RED
because in
tests,
light
OF
By
not
an
what
be
with
confined
we
illumination
means
the
considered.
by the leaf and transported
to
the
the
our
reactions
of
investigations
appearing
primary
Maybe
some
coleoptile,
in
to
the
primary
the indirect
the behaviour
leaf
influences
substance
maybe
the
is
of
the
formed
growth
of
10
G.
the
primary leaf is
that the
5.
influenced
substances
distributed
are
which
by the
light in such
orange
between
a
upwards from
transported
are
differently
and
coleoptile
manner
the
grain,
leaf.
primary
Summary
in
Orange light increases
the
probably perceived by
influenced by
the
7
of I.A. A.
application
be
light,
orange
of the
tip
(10“
primary
the
reduces
which
In
g/ml).
place of perception
this
the
of
OF
An
action
action
spectrum
lenghts
exercises
for
spectrum
obtained
by
and Owens
mesocotyl
the
caused
tip
of
light
orange
Avena
test
too
is
is
the unilateral
by
the primary leafproved
III
MOST
EFFECTIVE
LENGTH
influence
the Avena
inhibition
for
responsible
react
to
the
the
The
standard
which
of
of
light
is
coleoptile
growth in
the
the
longer
An
known.
not
mesocotyl
was
as
Weintraub and Price (1947)
well as by Goodwin
(1948). However, since it is uncertain whether the same
mechanism is
justified
for
on
The
light.
curvatures
THE
WAVE
wave
blue
light.
orange
CHAPTER
DETERMINATION
leaf.
too
case
size of the first positive
maximum
Avena coleoptiles the
induced by unilateral illumination with
curvature
to
BLAAUW—JANSEN
to
the way in which the
lengths
wave
larger
that the action
assume
than
spectrum
A
will
coleoptile
=
600 m/i,
be the
and
it
same
is
the
not
for both
reactions.
We
the
in
investigated which
phototropic
chapter
large
that
II. The
it
Coleoptiles
of Avena
impossible
light
it
is
of various
cultivated
irradiated
amount
wave
lengths
is
pre-illumination
applied).
of
The values
in
coleoptiles
influencing
described
the way
plant material however
determine
with
light which gives
of
these
given for
described
as
were
was
so
approximatively
an
(in
chapter
30-70
ergs/cm
2
a
30-70
ergs/cm
2
30-70 ergs/cm
2
30-70
ergs/cm
2
30-70
ergs/cm
2
30-70
ergs/cm
2
30-70
ergs/cm
2
30-70 ergs/cm
2
with
=
X
=
X
=
600
m
/I
=
620
m fi
=
640
X
X
under
the
shown
was
that the
from which
=
are
are
positive
m/i
m /«
/<
m/<
curvature.
in
Table VIII.
of six
experiments.
summarized
the
mean
VIII
Phototropic
X
580
it
of the direction
maximum
curvatures
560
II
II
unilaterally with
being pre-illuminated
they were
480
i.e. with the
ergs/cm 2 A
m/i,
TABLE
Pre-illuminated
chapter
after
20
experiments
the
in
prc-illuminated
independent
Immediately
one-sidedly
The results
to
most effective in
spectrum.
were
heading “phototropism”. They
effect of
was
of the
variability
proved
action
accurate
length
wave
curvature
curvature
in
13.1 ± 0.4
(109)
0.6
(51)
13.4
±
14.9
± 0.5
(64)
14.1
± 0.7
(36)
14.3
±
0.6
(51)
13.6
± 0.6
(46)
17.8
± 0.9
(36)
=
660
m/i
X
-
680
m/i
17.1 ±
1.3
(31)
A
=
700
m/u
12.6 ± 0,9
(36)
degrees
THE
INFLUENCE
In each
variability of
It
A
The
and
FAR
order
LIGHT
ON
difficulties
by
daily
the
light of the
lengths
wave
A
=
660 m/t and 680 m/t
decided
was
continue
to
with
experiments
our
of
variability
of
these
the
the
on
the increase in
reaction
phototropic
arrive at
to
To
experiments.
dependent
preferably
light
m/t.
means
complete
be
should
1922)
be
used
spectrum
more
process
used
indicator,
as
this
to
light
light,
blue
action
a
purpose
should
light
red
a
this
red
to
the
to
length of the coleoptiles. A sensitive
(Konxngsberger,
meter
lengths
wave
caused
joint effect of the variability shown by the response
the
directly
11
AVENA SEEDLING
THE
complete series of
a
avoid
to
however, makes it impossible
by
RED
plants.
the
So it
660
=
AND
effective in enhancing the maximum size of the first positive
curvature.
of
in
that
appears
most
was
RED
experiment the influence of
investigated
was
OF
auxano-
end.
Summary
In
chapter
this
X
lengths
of the
=
first
660
it
is
and
shown that with
680
is
m/t
positive phototropic
the Avena
caused
curvature
CHAPTER
THE
λ
=
by blue
of the
light
coleoptiles
enhancing
the
wave
maximum
size
light.
IV
OF THE AVENA COLEOPTILE TO LIGHT
REACTION
OF
in
effective
most
660
IN
mµ
CONNECTION WITH
ITS
GROWTH-SUBSTANCE METABOLISM
Data
1.
from
Several
red
the
authors
with
light
literature
have
their
tried
the
connect
too
different materials
they consider any attempt
cited
by them
were
discuss
to
sheer
as
reactions
metabolism.
(1957) summarize the literature
Galston
of the fact that
reports
to
growth-substance
on
this
their
own
plants
subject;, but
treated in
speculation.
of
Hillman
in view
different ways,
too
work
to
and
in
terms
We subscribe
of the
the view
to
of these authors.
Liverman and
Avena
red
coleoptile
and far
cylinders
red
Bonner
as
light
are
influence
In
heteroauxin.
to
(1953)
experimental
the
the
object.
only workers
According
choose
to
the
their results
to
the reactions of isolated coleoptile
hope that the influence of red light
the phototropic behaviour of the coleoptiles
might be explained
by the findings of these authors, we tried to reproduce their results;
on
without
however,
results
and
quantities.
ours
In
700
nated with
Liverman
our
be
ergs/cm 2
A
a
=
to
A
was
ours.
=
used).
Still
660 m/t
the
possibility
the
to
discrepancy
the
coleoptile
use
between
their
different
light
of
cylinders
whereas in
the
were
their
experiments
existed
that
heteroauxin
was
be
cannot
illumination
content
of
illumi-
experiments
larger quantity of light
light obtained from 2 daylight
Obviously
influences
the
660 m/t,
much
during 30 minutes (filtered
tubes
attributed
experiments
and Bonner
the
Probably
success.
should
fluorescent
compared
with
the
of
applied
light
of
coleoptile.
1
2
G.
To
the
avoid
of the
and
difficulties
determination
quantitative
coleoptiles
Maximal
BLAAUW—JANSEN
I.A.A.
estimated
was
by
growth of coleoptile
plants should
occur
differs
the
from
in
cylinders
2.
Cylinder
at
which
are
beset,
means
of the
I.A.A.
cylinder
derived
cylinders
and
extraction
the
content
test.
from illuminated
I.A.A. concentration of the medium which
an
I.A.A.
with
problems
of
concentration
maximum
inducing
growth
from non-illuminated
plants.
cut
(straight-growth
test
test)
METHOD
The
coleoptiles
I.
chapter
with
illuminated
darkness.
in
a
1
off
plants
700
and
of
of I.A.A.
a
solution
tions
on
the
a
The
sections
in
Van
and
sections
mm
floated
were
buffer of 0.0025
a
3
in
out
of A
light
green
length of the sections
All
sucrose.
=
tips
were
in
of
under
a
the
10 ml
adjusted
manipula-
After
560
measured
was
surface
in
placed
used
K-malcate
m
%
mm
in
was
kept
were
were
the
on
them
was
part
coleoptiles
Wey, and 3
der
3
next
described
as
part of
old,
660 mju,
=
pH of 4.5 and containing
carried
were
hours
A
vermiculite
hours
the illumination the
rejected.
Twelve
with KOH
2
microtome after
experiments.
88
were
ergs/cm
\ hours after
coleoptile
taken
the
in
cultivated
were
When
three
microscope.
EXPERIMENTS
A series
of
fluence of
a
experiments
range
irradiated and
of
the sections
units
3
=
shown
plants.
in
units
In
I
II
and
of sections
of irradiated
on
the
the in-
sections
final
micrometer
eye-piece
of
lengths
(50.9
of
plants
appear
show
confirmed in
Two
only
one
experiments
examples
are
In
to
be
10 -8
in
which
the
I
the
experiment
g/ml for
the
growth
however sections of
experiment II
show
whereas
concentrations,
to
In
ones.
to
experiments
8
against 2.10~ g/ml for the growth of
plants,
of non-irradiated
irradiated
used.
IX
estimated.
was
optimum I.A.A. concentration seemed
appear
tested
Table
of the
examples
are
concentration of I.A.A.
optimum
I.A.A.
each of which
was
mm).
Experiments
sections
in
out
concentrations
non-irradiated
are
carried
was
of I.A.A.
two
growth
sections
of
maximum.
growth
in which lower
presented
maxima
at
different
non-irradiated
This
I.A.A.
plants
phenomenon
concentrations
III
in Table IX,
and
was
were
IV and
in
Fig. 3.
DISCUSSION
Pre-illumination
of
doubtedly influences
react
on
growth
I.A.A.
curves
non-irradiated
1.
shows
The
two
intact
The essential
of
coleoptile
plants may be
seen
whereas
in
of
between
obtained
two
curve
the
light
A
=
660 m/t
un-
coleoptile sections subsequently
difference
sections
concentration-growth
maxima,
with
plants
the way in which
the concentration-
from
irradiated
and
fundamentally different ways.
of irradiated
curve
of
coleoptile
non-irradiated
sections
sections
INFLUENCE
THE
OF
AND
RED
FAR
RED
TABLE
lilumination
with
700
Concentr.
of
2
ergs/cm
A
=
LIGHT
THE
ON
13
AVENA SEEDLING
IX
660
m,u.
Final
length
of
sections
I
II
Plants
Plants
I.A.A.
in
g/ml
illuminated
0
not
52.0 ± 0.7
6.10
s*
53.0 ±
—
8
i.io-
54.1
8
2.10-
53.7
6.10-8
±
illuminated
illuminated
0.4
48.5
—
0.3
51.1
± 0.4
± 0.2
51.4 ±
0.4
54.4 ± 0.3
53.4 ±
0.2
50.6
53.2
54.9 ± 0.3
—
not illuminated
0.5
±
± 0.2
—
—
± 0.3
—
50.1 ± 0.4
55.1
± 0.3
52.1 ± 0.3
55.1
±
0.5
54.3
55.8
±
0.6
7
53.3
±
e
52.8
± 0.5
52.5 ± 0.5
I.IO-
51.2
± 0.3
51.0
± 0.2
0
48.8
±
0.4
51.4
±
0.6
50.6
±
I.IO-10
52.5
± 0.3
50.0
±
0.3
51.2
± 0.3
51.9
± 0.3
1.10-9
51.9
± 0.4
50.2 ± 0.2
52.6
± 0.1
52.6
± 0.3
9
55.8
±
0.5
50.5
± 0.2
54.1
±
0.3
51.9
8
50.5
±
0.3
53.5
± 0.3
52.3
± 0.2
54.3
±
0.3
55.4
±
0.5
55.5
±
0.3
54.7
±
0.3
55.8
±
0.3
55.9
± 0.2
56.5
±
0.5
55.8 +
0.5
I.IO1.10
5
0.5
53.3
± 0.3
0.5
±
—
—
III
3.10i.io3.10-
8
I.IO“7
Fig.
3.
Final
length
reached
non-irradiated seedlings
shows
only
enon
that
one.
very
and
In this
low
IV
in
I.A.A.
from
case
no
carried
this
to
2.
Hancock and
out
the
The
Both show
diated
two
two
sections
(1957)
of wheat
of
I.A.A.
phenomenon
Lacey
with sections
displacement
of
an
as
has
coleoptiles.
endogenous
concentration-growth
to
be
found
at
an
paid
also
curves
I.A.A.
to
in
by
are
from
cut
660
m
/<.
the
the
been
=
phenomgrowth of
reported
tests
The authors
auxin
maxima, but whereas the first
is
sections
inhibit
appearing
0.2
±
53.0 ± 0.2
coleoptile
attention has been
0.2
52.4 ±
seedlings irradiated with light of A
concentrations
non-irradiatcd sections. This
Barlow,
solutions by
0.1
that
by
were
attribute
I.A.A.
essentially
equal.
maximum of the irraconcentration
of about
14
G.
9
3.10
g/ml, the first maximum for the non-irradiated sections appears
I.A.A.
an
sections
concentration 0. So
is
regarded
concentrations
In both
blank
It
as
cases
was
hoped
to
The
did
curvature
shifted
at
for the irradiated
curve
I.A.A.-
higher
to
for the non-irradiated sections.
but
in
the
show
possibility
the
and
curve
grow,
would
curve
the
case
the fact that sometimes the sections in the
not
whether
investigated
exclude
this
the
to
over
pass
buffer solution
in
compressed
as
compared
we
centration-growth
3.
BLAAUW—JANSEN
that
did shrink
our
test
conclusion
contrary.
the
too
in
maxima;
two
the
on
curvature
this
we
way
due
was
con-
bias.
to
test
METHOD
The
the
plants
test
When the
with
plants
2
70
described
as
in
II
chapter
under
whereafter
after
(compare
the
prevent
others
about
LA.A.
was
and
applied
six
an
the
illuminated
were
in
were
carried
out
in
primary leaf
induce
two
from
block
agar
light.
orange
loosened
was
was
to
maxima of
consuming
maximum
(at
with
substance
low
light
of
So in order
curvature.
the
a
unilaterally.
applied
whether irradiation
investigate
to
first
the
decapitation
on
was
decapitation
coleoptile
brings
second
decapitation
II),
was
660 m/u would
tested
old, part of them
The
light from influencing the reactions of the coleoptile
chapter
intention
660 m/t.
=
the
the orange
prevent
=
A
first
the
Immediately
Our
88 hours
were
darkness.
ergs/cm
kept
11 hours after the irradiation the plants remained in darkness,
During
A
cultivated
were
heading “Standard Avena-test”.
that
to
possibly
I.A.A.concentration)
immediately after the first decapitation, and the
was
omitted.
Each
concentration
of I.A.A.
was
coleoptiles.
EXPERIMENTS
The
size
of
the
experiments
two
Table X.
Fig.
In
curvatures
hours after
4
the results
they
as
the
of
of
experiment
II
TABLE
Curvature
in
the
I
Concent,
of
Avena
test
after
in
are
different
three
I.A.A.,
are
given in
plotted.
X
illumination
II
plants
appeared
application
with
700
ergs/cm
2
III
plants
A
=
660
m/j.
plants
I.A.A.
not
in
g/ml
illumin.
illumin.
IO"
0.4
10- 7
0.5
10-
7
13 ± 2
10-
7
15 ± 2
0.7
10- 7
0.8
10-
0.9
10- 7
7
1
19
±
3
1
6
±2
8 ±
1
±
1
7
±2
9 ±
I
13 ± 2
17
± 2
11
± 2
16
±2
13 ±
1
15 ± 2
18 ± 2
13
±2
18
± 2
11 ±
2
16
15
± 3
20 ± 2
18 ±2
18 ± 3
18
±
1
17 ± 2
24
± 2
± 2
18 ± 2
22
±
I
21
±2
io-
25 ± 3
27 ± 2
21 ± 2
21
±
1
2.0
IO“7
26 ±2
30 ± 2
27 ± 3
23
±
1
4.0
10-7
30
± 3
32
4-
1
30 ± 2
23
±
4
6.0
10-
7
33
± 3
33 ±
1
1.0
7
illumin.
±
—
23 ±2
not
illumin.
6
—
24 ±
illumin.
11
7
0.3
0.6
not
illumin.
10 ± 2
11
14
±2
± 2
17 ±
1
20
±
1
24 ± 2
19
±
3
24 ± 2
27 ± 3
30
±
5
25
±
1
28 ±
1
INFLUENCE
THE
OF
RED
AND
FAR
RED
LIGHT
ON
THE
AVENA
15
SEEDLING
DISCUSSION
On
condition
that
each
curvature-concentration
plants,
irradiated
irradiated
the
or
4.
Curvature
and
the
three
I.A.A.
than
at
to
shown
average
the
not
as
above-mentioned
solution
were
curvatures.
curvature
nearly
plants.
so
the
as
III
curvature
0.7
the results
from
I
experiments
curvature
or
The first
ones.
X
as
test
by
the
10~
7
X
—
the
irradiated
plants
g/ml is significantly
ones.
being
day
might be
studied.
irradiated
due
were
the
to
each
same
series.
I.A.A.
the
between
the
Table
XI
plants
of series
not
observed
of
of three series of
treated
the
in
exactly
series
shows
a,
is
variability
non-irradiated
concentration
an
large.
too
to
at
the varia-
curvature
curvatures
They
and
more
larger
it
Presumably
day
the
a
m/x.
experiments,
irradiation,
were
the
660
of the three
to
as
In each of
non-irradiated coieoptiles
light of
of
II
platform.
a
the
non-
maximum of
and
experiments. Agar blocks soaked in the
observed
large
to
applied
At
the
experiment
the
the
plants
way
same
in
I.A.A. concentration
non-irradiated
itself,
for
that the differences in
argument
the
plants and
by
maximum
irradiated with
by the plants
same
considered
of the non-irradiated
curvature
counter
the
the
in
coieoptiles
experiments
the
To
in
concentration of 0.6
admissible
bility
reached
by
shows
the irradiated
plants appears
pronounced peak,
less
Fig.
for
two
is
one
experiment
curve
in
same
the
resulting
differences
b, and
the
I.A.A.
c
are
in
not
those observed between irradiated and non-irradiated
16
O.
BI.AAUW—JANSEN
TABLE
I.A.A.
at
a
series
7
this
and
11
± 2
8
±
I
15
±
1
11
±
2
12
±
0.7
IO-
13
±
1
11
±
I
12
± 2
0.8
10-7
15
±
0
15
±
1
16
± 2
0.9
10-7
16
± 2
16
±
1
15 ± 2
16
± 3
17
± 2
16
3
16
± 2
18
± 2
7
1.0
IO“
18
±
1.2
IO-7
19
± 2
17
±
2
17
±
1
1.8
10- 7
19
± 2
20
±
1
20
±
1
19
±
± 2
17
± 2
19
±
7
1.4
10-7
1.5
IO“7
case
first
21
it
too
18
20
possible
is
that
plants
is
possibilities
the
that
irradiation
with
the
plants
hypothesis
is lower
the
on
of A
light
than
possibilities
the
660
=
m /.<
the first maximum.
causes
with
correspond
discussion
a
straight-growth
content
The results
of such
results
or
extraction
compound,
new
necessary.
Summary
I.A.A.
endogenous
of
4.
2
and
(par.
test
1
2,
and
an
in
=
the
quantitative
will
analysis
660 m/i
enhanced
be
analysis
given in
chapter.
been
the
or
shown
that
irradiation with light
activity-concentration
the
of
curvature
additional
peak
is
curve
coleoptiles
situated
at
a
QUANTITATIVE
IRRADIATED
purpose
content
ANALYSIS
AND
of this
of the
of
X
=
660
is
plotted against
low
I.A.A.
part
causes
OF
the
of the
an
growth
additional
of coleoptile
I.A.A. concentration.
concentration.
V
THE
I.A.A.
NON-IRRADIATED
coleoptiles.
m/i
which is obtained if the
CHAPTER
I.A.A.
between irradiated
investigate whether irradiation with light of A
to
the
next
The
applied in these experiments.
synthesis of a substance which
in
also
plants
should be situated
then this
of irradiated
content
2
12—13).
the
I.A.A.
non-irradiated
1
±
ones.
possibility
two
would be
sections
was
1
might be accounted for by
I.A.A.
endogenous
synthesis
in
the
but
±
the difference in behaviour observed
case
In order
This
1
± 2
maximum of curvature,
second
has
1
1.1
These
peak
9
10-7
considered
It
± 2
10~
stimulates the
lowers
c
IO'7
that of non-irradiated
p.
series
0.5
that the
see
degrees
b
0.6
non-irradiated
A
in
series
a
lower I.A.A. concentration than
a
In
g/ml
this
In
Curvature
concentration
in
have
XI
investigation
Destruction
as
CONTENT
OF
COLEOPTILES
was
well
to
as
estimate
the
formation of
during extraction
tions caused
should be prevented;
moreover, complicaby the formation of auxin complexes should be avoided.
The extraction
described
by
method
Terpstra
used
in
(1953).
these
experiments
is
based
on
that
THE
INFLUENCE
OF
AND
RED
FAR
LIGHT
RED
THE
ON
AVENA
17
SEEDLING
METHOD
Coleoptile
quartz sand;
tips
of
this
was
was
mixed
and
kept overnight
10
adding about
ml of
10
The
10
aqueous
residue
dissolved
This solution
in
a
applied
was
of Whatman
No.
The
chromatograms
about
18 hours.
A marker
After
into
drying,
and
sprayed
a
The
of ether
and
cut
from
for 5
agar
3
mm
slices
in the
were
the
with
the residue
chromatogram.
a
solvent
was
of
employed
was
isopropyl
for
with
the
parallel
run
solvent.
as
ascending
strip of paper with
with
1
a
dried
was
solution
°/ 0
the marker
of
with
the
at
37° C.
and the
residue
into 6 blocks
of the
position
and extracted
The ether
as
by
fractions
Terpstra
and
tested
a
yellow-
0.6.
I.A.A.-marker
with 3
an
10 ml
X
combined
were
transferred into
was
described
cinnamic
again and, when brought
ranged between 0.4 and
chromatogram,
manner
cut
and
HCl-gas, showed the marker I.A.A.
minutes
evaporated,
of 20
strip
corresponding
zone
three times
mixture
a
with the
the
The Rf-value of I.A.A.
was
spot
methanol. The
container with
red spot.
HG1, methyl orange
extract.
removed
in
1: 1)
of I.A.A.
spot
and
aldehyde
with
of ether.
and
used,
(8:
N
line of
starting
water
developing
was
spot
of
by
discarded.
was
evaporated,
amount
developed
were
chromatogram of the
the
were
alcohol, 28 % ammonia and
C,
was
ml
three times
and shaken
were
small
to
1
5
another
This ether fraction
(final pH ca4),
more
mass
made alkaline
was
shaken
was
about ■—5°
.
These ether fractions
once
at
some
ground tissue
day the
with
acidified with 0.1
was
indicator
as
10 ml of ether.
Sheets
g
rinsed
was
Next it
NaHGO
mg
next
thus obtained
ether, freed of peroxides.
being used
was
tissue
extract
The
C, frozen
The
temperature.
The
ml of
100°
with
ground
were
light.
orange
water at
tap
this
at
filtered.
The
length
mm
performed in
with 5 ml of
thawed and
tapwater.
6
about
agar
slice
(1953). Next day the
on
six
colcoptiles.
EXPERIMENTS
The I. A.A.-content of
by
1|
means
hours
of
the
ergs/cm
are
given.
2
).
tips
the
before
700
tips of non-irradiated coleoptiles
curvature
660
—
m,«.
with
test
that
collected
were
In Table XII
TABLE
of
had
tips
extract
15
compared
from
± 2
10
13 ± 3
30
29 ± 3
in
degrees
from
extract
with
irradiated
A
=
660
6
±
1
8 ±
1
15 ± 2
10
34
3
12 ± 2
36
2
17 ± 2
±
light
m,u
34 ±
± 3
tips
non-
10
±
with
experiments
10
40
which
irradiated
XII
irradiated tips
50
been
the results of six
Curvature
Number of extracted
was
tips of plants,
1
of
G.
18
In
the
lowers
half the
one
irradiated
6
test
been
1-| hours
the
original
it
tips
irradiation
I.A.A.
For
content.
appears
with
of the
content
extract
to
the various
I.A.A.
necessary
experiments
of the
content
X
of Terpstra
XII,
of Table
extracted
tips
from
tips
each
to
It
has
not
curvatures
differences
to
differences
to
m/t
about
I.A.A.
of the
due
660
to
(1953).
are
or
=
tips
10
about
investigated whether the differences in the size
the
of
light
coleoptile
the determination of
This matches the results
plants.
found in
in
of
course
doubtlessly
BLAAUW—JANSEN
in
the
sensitivity of the testplants.
These
that
results
than from
the
confirm
auxin
more
of
statement
be obtained from
can
Van
Overbeek
(1936),
tips of non-irradiated coleoptiles
tips of coleoptiles irradiated with orange light.
Summary
with
Irradiation of Avena seedlings
the
I.A.A. content
half
the
of the
found
content
of the
tips
in
the
dark
Further
for
the
OF
the
coleoptile
a
the absence
the
of
non-irradiated
first
adding
=
1J
660
lowers
m/i
hours
to
about
desirable
make
to
irradiation
the
in
peak
should
be
(R.L.F.)
whether
sure
with
If this
red
growth-concentration
due
to
ihe
fact
light
would be
so,
of
curve
that the hetero-
of these coleoptiles is too high to allow the development
maximum, and then it would be impossible to introduce
into
peak
the
latter
larger
larger
the
similar
extract
case
effect.
would
of
the
called
“Red
unless
ones,
surface
depend
extract
would
first
I.A.A. concentrations when
henceforth
a
non-irradiated
the
on
ratio
supplied
be
growth
larger
Light
by
coleoptiles
I.A.A.
would
be
which it had been adsorbed.
to
dominate, and then
of I.A.A.
Thus
the
irradiated
fr'm
amount
amount
of
curve
would
it
two
the
the
from
extract
an
which of these
the
A
of
FACTOR”
decrease of the I.A.A.-content.
coleoptiles
displaced by the
In
LIGHT
of the
consequence
first
2
course
content
of the first
this
ergs/cm
the
VI
“RED
seemed
investigations
only consists in
auxin
THE
in
controls.
CHAPTER
EXTRACTION
700
light of
coleoptiles
is
to
that
the
required
to
shift
of the extracted
or
extract
coleoptiles,
should
peak
amounts
I.A.A.
should expect
the
to
that
Factor”
we
“R.L.F.”,
obtain
to
a
higher
compound,
are
supplied.
METHOD
Extracts from
as
described in
bio-assayed,
contains
the
the
in
solutions
in
accordance
effect
were
of
the
However,
was
as
were
before
indicated
by
prepared and purified
the
an
discarded. The eluates
transferred into agar blocks
of I.A.A.
with
V.
which,
zone
I.A.A.,
chromatogram
the
tips of irradiated seedlings
chapter
of various
the
results
‘R.L.F.’
is
the
from
as
preceding
observable
only
the
rest
was
spot,
of the
that had been soaked
concentrations
of
chromatogram
I.A.A.-marker
it
was
supposed,
experiments,
when it
is
that
applied
in
THE
INFLUENCE
OF
AND
RED
combination with I.A.A.
in
II.
chapter
They
and
decapitation,
RED
FAR
LIGHT
ON
19
AVENA SEEDLING
THE
The test
plants were cultivated as described
kept in darkness till the very moment of
were
treated
described in
as
chapter V.
EXPERIMENTS
First
from
cm
2
chromatogram obtained with
a
that
plants
?.
=
660
divided
was
m
tested
in
tested
eluted and
was
Rf-values
lower
combination
with
than
with
Eluate
that
that
of
of
of
cut
ergs/
starting
line,
contained all
eluate
I.A.A.,
Both
\
tips
700
chromatogram
the
a
LA.A.
I.A.A.-solutions
with
The
activity.
bio-assayed.
than
hours
1|
parallel
cut
a
higher
Rf-values
for
its
on
by
from 300
extract
an
irradiated
halves
two
with
with
ones
been
was
fi,
into
and each half
substances
had
eluates
10~ 7 and
X
f
the
b
the
were
10~ 7
X
g/ml.
TABLE
XIII
in
Curvature
I.A.A.
concentration
in
Control
I.A.A.
g/ml
with
with
a
combined
eluate b
0.50
X
10-7
11
±
1
10
±
1
11 ±
1
x
10~7
11
±
1
4
±
1
11
I
Table
in which
XIII,
The
experiment
the
Only
part
eluted, but it
the
more
the
first
line.
starting
of the
curvatures
than
plants
test
±
given,
are
Possibly it contained
an
optimum concentration.
with
repeated
was
of
was
the
inhibited the curvature.
a
active substance in
0.56,
eluate
I.A.A.
0.75
shows that eluate
to
degrees
combined
divided into 6
The Rf-value
made from
extract
an
above
chromatogram
of 1.4
zones
in
of I.A.A.
the flow distance of the solvent 22.5
the
cm.
I.A.A.
by
cm
this
15
place
cuts
was
parallel
experiment
The result
tips.
was
is presented
Table XIV.
in
TABLE
I.A.A.
concentration in
10~
1
X
10~7
+
zone
1
X
10“ 7
-j-
zone
2
10
degrees
10 ±
15 ± 2
-7
(front)
X
10
zone
3
1
X
10“
-j-
zone
4
7
1
X
10“7
+
zone
5
22
X
10 -7
+
6
8
+
7
The eluate
the
zone
from
zone
curvature
5,
1
10 ±
1
enhances
with
slightly
induced
1
1
±
±4
±
1
higher Rf-value than I.A.A.
I.A.A.
by
± 2
(IQ-
7
g/ml)
from
about
22°.
Now
it
found for
an
in
X
1
to
Curvature
g/ml
7
1
1
15°
XIV
eluate
was
tried
irradiated
from
a
to
imitate
plants
chromatogram
To this purpose extracts from
been irradiated
hours
the
curvature-concentration
by supplying
in
non-irradiated
combination
eoleoptile tips
previously
with 700
cut
curve
plants
with
with LA.A.
from
ergs/cm
2
X
plants that had
=
660 m/u,
were
20
G.
freed
from
I.A.A.
determined
was
above this
zone
The
by chromatography.
by
of
means
marker
a
extracted
was
of the eluate
amounts
BLAAUW—JANSEN
and
The
bio-assayed
as
combined with various
were
I.A.A.
containing
zone
spot.
a
chromatogram
whole.
Various
of
amounts
I.A.A.,
and the mixtures tested.
results
The
of three
II
Experiments
typical
III
and
experiments
plotted
are
TABLE
in
given
are
Fig.
in Table
XV.
5.
XV
I
EXP.
Amount
I.A.A.
of
added
extract
in
g/ml
from
extract
none
4/6 tip
extract
from
8/6
i
±
i
4
±
1
6
±
1
9
±
1
13
±
1
16
±
3
8
± 2
10
±
1
9
±
1
13
± 2
14
± 2
9
± 2
13 ± 2
15
± 2
17
±
17
16
± 2
17
±2
X
IO“
1.00
X
IO“ 7
1.20
x
IO“
7
10- 7
1.40
x
1.60
x
10-
1.80
x
10- 7
7
1
±
NO Ö o
X
o ij
19 ±
1
o
X
o
il
25 ±
0
in
plant expressed
test
tips
7
0.50
O
per
coleoptile
concentration
tip
3
n
EXP.
Amount
I.A.A.
of
added
extract
in
g/ml
X
10- 7
0.62
X
10- 7
fr.
extr.
none
0.50
9
dr 2
9
±
19
dr
4
± 2
12
±2
10
dr
13
dr 2
15
dr 3
17
dr 2
20 db
10-
7
10 ±
1
x
10-
7
13
1.00
x
10- 7
1.25
x
10- 7
EXP.
fr. 4/6 tip
extr.
1
14
0.87
2/6 tip
dr
10
7
1
x
in
plant expressed
test
±4
7 ± 2
±
0.75
per
coleoptile tips
concentration
extr.
fr.
9
8/6 tip
± 2
1
10
d:
1
16 dr 2
11
dr
1
3
5
±
1
11
dr 2
12
dr
2
15
dr 2
13
dr
3
4
Ill
Amount
I.A.A.
added
of extract
in
g/ml
extr.
none
fr.
2/6 tip
extr.
in
plant expressed
test
tips
fr. 4/6 tip
extr.
fr.
8/6 tip
1.00
X
5
±
1
6
±
1
5
±2
6
±
1
1.25
x
10- 7
6
±
1
7
±
1
4
±
I
5
±
1
1.50
x
IO- 7
9
± 2
9 ± 2
8
± 3
8
±
1
1.75
x
10~7
15
± 2
4
15
±2
2.00
x
10-7
19
±
± 2
x
IO-7
17
x
IO- 7
18
2.25
2.50
It
IO“
7
per
coleoptile
concentration
appears
transforms
irradiated
amount
obtain
a
These
that
the
a
15
±
1
14
±
3
11
± 2
10
± 2
9
± 3
17
± 3
12
±
1
12
± 3
14
±
purified
extract
one-peaked
coleoptiles
of eluate
into
16 ±2
1
from
two-peaked
equivalent with the
1
tips of irradiated coleoptiles
curvature-concentration
a
±
15 ± 3
extract
one.
from
curve
Application
1
of
non-
of
tip is sufficient
an
to
maximum effect.
experiments
of non-irradiated
test
show
plants
that
to
the
R.L.F.
growth
can
affect
the response
substance. The I.A.A.-content
THE
Fig.
5.
INFLUENCE
Influence
light
Fig.
6.
on
RED
which
exercises
Influence
light exercises
OF
which
the
Right:
a
on
a
AND
purified
the
plants
RED
extract
curvature
purified
curvature
Test
FAR
of
from
of
extract
test
LIGHT
THE
AVENA SEEDLING
coleoptiles
non-irradiated
from
plants.
irradiated
ON
with
irradiated with
21
red
coleoptiles.
coleoptiles irradiated with red
Test plants not irradiated.
Left:
light
of
A
=
660
m/i.
22
G.
of these
therefore is
plants
BLAAUW—JANSEN
not
high
so
shifts
I.A.A.
higher
to
Either
supplied.
are
concentrations
this
is
not
so,
to
as
the R.L.F. No evidence is found for the
if
or
make
them insensitive
that the first
supposition
larger
the shift
of R.L.F.
amounts
is
too
to
peak
small
be
to
perceptible.
Finally the R.L.F.
and
was
non-irradiated
in the
plotted
tested in the
Table
plants.
graphs
6
Amount of
XVI,
in
plant
Not
10-
7
0.75
none
1.00
none
1.25
X
IO“
4/6 tip
0.75
x
4/6 tip
1.00
x
10-
4/6
tip
1.25
x
10-
8/6
tip
0.75
x
10" 7
8/6
tip
1.00
x
10-
8/6
tip
1.25
x
10- 7
are
XVI
Pre-illuminated
pre-illuminated
X
16
±
3
19
±
1
16
±
±2
14
±
1
10- 7
24 ±2
30
±
3
7
18 ± 2
19
±
2
18 ± 3
16
± 3
io-
X
7
7
21
7
7
27 ±
3
I
21 ±
3
19
± 2
17 ±
2
16
±
3
19 ± 2
23
±
2
II
Amount
of
I.A.A.
added
concentration
in
plant
test
Not
Pre-illuminated
pre-illuminated
g/ml
7
none
0.50
X
10~
none
0.75
x
10- 7
8
7
± 2
8
11 ± 2
17
1.00
X
10^
2/6
tip
0.50
x
10-
2/6
tip
0.75
2/6
tip
1.00
x
10-7
19 ± 2
4/6
tip
0.50
x
10- 7
12
±
1
4/6
tip
0.75
x
10-
7
17
±
1
4/6
tip
1.00
x
10- 7
17
±
4
10-
7
0
±
0
3
7
18
2
25
none
8/6
10
x
0.50
tip
x
tip
0.75
x
10-
8/6
tip
1.00
x
io-
In
6
Fig.
reproduced
Fig.
7
7
15
7
with
together
(based
on
and non-irradiated
added,
2/6 tip,
in
either
plants,
in
with
the
graphs
26
±
10
±
1
2
17
±
3
±
±
13 ± 2
21
testplants
7b
are
after
±
2
i
1
± 2
20 ± 2
the
I)
R.L.F.
the
graphs
or
not
irradiated
the
of
the
(a),
plants
are
(b).
graphs of irradiated
plotted together, in Fig.
addition
± 2
17
with
of
3
19 ± 3
experiment II) however,
Fig.
2
± 3
2
experiment
supplied
±
± 2
±
16 ± 3
(corresponding
R.L.F.
These
26
7
8/6
non-irradiated
In
which
g/ml
none
extract
per
irradiated
on
of
I.A.A.
concentration
added
test
EXP.
figures
I
extract
per
experiment
the
and 7, shows the results.
TABLE
EXP.
same
7a with
of the R.L.F.-extract
no
from
Fig. 7c from 4/6 tip and in Fig. 7d from 8/6 tip.
experiments indicate that the R.L.F. enhances the curvature
of non-irradiated and irradiated
concentration,
independent
plants
of the
at
the
amount
same
exogenous
of R.L.F.
I.A.A.-
INFLUENCE
THE
RED
OF
AND
FAR
RED
LIGHT
THE
ON
23
AVENA SEEDLING
DISCUSSION
It
impossible
appears
the
in
plants
test
to
their
to
attribute the absence of the first maximum
size of the
the
presenting
curve
This appears from the fact that,
7.
Fig.
Influence
exercises
light
plant supplied
from
can
of
with
4/6
irradiated
purified
a
d\
coleoptile
and
test
tip.
c:
Each
possibilities
for
light of
with
660
from
extract
b
with
8/6
first
a
Each
:
red
test
extract
tip.
maximum
the different behaviour
explaining
=
from
irradiated with
plant supplied
test
coleoptiles
X
extract
added,
extract
development of such
non-irradiated
irradiation with
No
a\
plant supplied
the
tips,
plants.
purified
coleoptiles
from
extract
a
are
left:
m/t may
cause
or
increase
formation of R.L.F.
Secondly,
capacity
As
the
an
irradiation
of the
with
coleoptile
argument in support
observation
that
light
react
to
“dark”
right
the
were
ones,
in order
that
to
in
of this
reach
required
fulfilled
coleoptiles
supply
to
our
—
m/u may
already
explanation
coleoptiles
to
660
R.L.F.
can
react
enhance
the
present.
we
to
may
conceive if the second
adduce
supply
a
of
explanation
one.
If irradiated
the
of X
to
of the first
R.L.F., which would be difficult
due
I.A.A.-level.
by applying
test
from 2/6
Each
be obtained. Two
Firstly,
of
curvature
extract
tip.
irradiated
the
which
the
on
of non-irradiated
curvatures
high endogenous
too
by
the
contain
substance
“optimum”
range
but
R.L.F.
required
than
this
ones.
non-irradiated
by irradiated coleoptiles
concentration,
the non-irradiated
experiments,
the limited
more
This
would be
expectation
negative result
of concentrations that
were
may
less
than
was
possibly
tested.
not
be
24
G.
BLAAUW—JANSEN
SUMMARY
to
660 m/t it appeared possible
irradiating seedlings with light of X
the
coleoptile tips a compound (R.L.F.) which causes in the curve
After
=
isolate from
of
similar
a
that
of
irradiated
in
this
already
in
irradiated
peak
and
in
test
I.
Discussion
With
be
can
1.
regard
identity
curvature-concentration
of
amount
in
curve
the
at
R.L.F.
same
supplied
VII
LIGHT
RED
of
the
of
appearance
naturally
plants).
the
of the
identity
the
to
the
occurs
FACTOR
r.l.f.
R.L.F.
only
indications
vague
given in this chapter.
It
that
the
on
THE
as
non-irradiated coleoptiles
CHAPTER
IDENTITY OF
the
independent
irradiated
the
(in
present
concentrations)
I.A.A.
first
concentration,
non-irradiated coleoptiles
in
low
The
ones.
case
I.A.A.
exogenous
or
maximum (at
first
the
appears
of curvature
the degree
presenting
it
is
probably
be
can
acid.
an
transferred
This
by
can
shaking
be
the
concluded from
from
acidified
an
fact
aqueous
solution into ether.
2.
In
hours
R.L.F.
is
alkali.
stra
in which
experiments
either
in
acid
The
1.5NHC1
the R.L.F.
in
or
heated
was
1.5NKOH, it
proof, but that it looses its activity
experiment
The
(1953).
the R.L.F. either
made
was
untreated
according
induced
curvatures
if it
treated with acid
or
two
that
boiled
is
the
with
the method of Terp-
to
mixtures
by
during
appeared
of
and
I.A.A.
alkali,
or
are
given
in Table XVII.
TABLE
Curvature
I.A.A.
concentration
in
1
0.75
x
10“
21 ± 2
1.00
x
10-7
28
±
29
± 2
The
7
7
10-
x
were
of chapter
that
the
possible
polluted
it
causes
R.L.F.
in
only
with
additive
activity
with
boiled
alkali
3
1
8
± 2
27 ±
±
1
16
±2
28 ± 2
19 ±
I
3
20 ±
were
30
applied in this
±
series of
in
the
of
plant
±
the
respond
an
of
For
3
experi-
experiments
this
and
is
without addition
it
reason
auxin-a.
Kogl’s auxin
of
that
might
auxin-a, but
as
is
improbable
This
preparation
I.A.A.
preparation
his
be
the
would
had
be
been
curvatures
ascribed
preparation
to
an
was
a
hardly conceivable.
(1957
gibbercllic acid
Gibbercllic acid
coleoptiles,
this
Kogl’s
amount
this
of I.A.A.
to
Avena
with
that
event
large
one,
to
curvatures.
Brian and Hemming
solution.
boiled with acid
addition
test
per
8
that
identical
solutions
highly purified
to
tip
one
14
applied
no
is
so
by
4.
the
after
degrees
from
VI.
induced
fail
in
higher than these used
considerably
If the R.L.F. is
of I.A.A.
untreated
3
of R.L.F.
amounts
ments,
3.
extract
Control
g/ml
1.25
XVII
and
1958)
if there is
enhanced the
found that
no
I.A.A.
pea
in
epicotyls
the culture
growth-promoting activity
of
INFLUENCE
THE
I.A.A.
at
all
the
OF
RED
I.A.A.
conformity with the
in
gibberellic
Moreover
acid
in the
4
/tg
chromatogram
dried
again
60° C
acid
The I.A.A.
detected
So
nearer
to
experiments
and
1.5
purpose.
methanol, and
4
treatment
the spot
light.
seen
Table XVIII.
in
XVIII
Rf-values
0.59
0.34
0.54
0.31
0.48
0.62
0.74
the Rf-value
Rf-value of
that
of
gibberellic
X
this
the
developed,
in
for:
acid
R.L.F.
0.52
to
of
0.45
>
acid
gibberellic
be
like
found,
-8
agar
blocks
coleoptiles. The coleoptiles
and
chapter V,
were
were
Preliminary
acid
the
ranges from
0.5
R.L.F., somewhat
with
advance
acid.
gibberellic
test
soaked in mixtures of I.A.A.
showed
on
10~ 8
that
decapitated
once
darkness
and
g/ml
suitable for
cultivated and tested
absolute
in
exclude
not
concentrations
tested
were
kept in
tests
were
were
does
identical
curvature
acid;
agar slices
g/ml gibberellic
The
is
R.L.F.
gibberellic
acid.
acid
gibberellic
the
these experiments
10
be
can
Gibbcrellic
compound is
possibility
For
2
chromato-
the front than I.A.A.
Activity
3.
and
parallel strip of the chromatogram
0.49
that
this
Thus the
the
gibberellic
test.
being
H S0
%
1
U.V.
on a
0.35
appears
curvature
After
with
in
seen
I.A.A.
0.7.
not
gibberellic acid
V.
TABLE
It
the
minutes. After
10
The
to
tested. This is
were
chapter V.
cf 5
The results
25
SEEDLING
proof.
of
chapter
be
AVENA
with 4 fig I.A.A.
mixed
was
during
was
spot
described in
as
and in
Rf-value
in
can
that
THE
investigate the activity of gibberellic
to
dried, sprayed
was
at
gibberellic
of
acid
ON
determine the Rf-value of
to
test
the
described
as
LIGHT
acid
not
while
straight-growth
gibberellic
graphed
is
acid
of
RED
in which the R.L.F. interacts with I.A.A.
chromatograms, and
our
Determination
2.
FAR
concentrations
way
Yet it seemed worth
acid in
AND
till
as
our
Avena
described in
the
moment
two
hours
of
decapitation.
Table XIX shows the
of the
application
ment II
is
plotted
Obviously
an
4.
agar
in
Fig.
gibberellic
additional
curvatures
blocks;
8
acid
two
of the
test
plants
experiments
were
test
after
Experi-
(left).
can
induce
at
low I.A.A.
peak in the curvature-concentration
Straight-growth
made.
with
i.a.a.
combined
concentrations
curve.
with
gibberellic
acid
Finally
some
I.A.A. and of
cylinder
tests
were
carried
out
with mixed
solutions of
gibberellic acid. The method is described in chapter III.
26
G.
BLAAUW—JANSEN
TABLE
EXP.
g/ml
degrees
gibberellic
io-
0.
0.6
x
10- 7
0.7
x
U)-
7
0.8
x
10-
7
0.9
x
10-
7
7
1.0
x
10
1.1
X
io-
1.2
x
10
7
7
x
10~7
0.62
X
10
0.75
x
10-
EXP.
acid
in
6
g/ml
1.5
6
10
x
12 ±2
11
±2
11
H
±
11
± 3
13 ± 3
14
±
23
±
1
2
20
1
26
±
1
23
±
2
23
±
24 ±
15
2
17
±
1
± 2
15
±
0
±
17 ± 2
23
±2
21 ± 3
±
2
22+3
1
23 ±
2
26 ± 2
1
25 ± 2
26
±2
II
7
5
±
1
9
7
±0
4
±2
±2
8
±
1
4
±
t
2
-
10
7
9
10
± 2
7
± 2
x
io- 7
12
±
1
19
± 2
7
±
1.12
x
10-
13
± 2
13
± 2
7
±2
1.25
x
7
10
± 2
7
6
±
2
1.37
x
10- 7
12
±
2
19
16
±
1
1.00
Fig.
7
x
0.87
8.
7
10
Influence of
cylinder
gibberellic
dots:
test;
no
acid.
The
coleoptiles
that
were
pre-illuminated with
In
Table XX the
hours
scale
they
units;
8
floated
two
(right).
x
10-
be
to
red
or
in
Left:
added;
G.A.
1.5
Fig.
of
concentration
I
1.3
not
in
Curvature
I.A.A.
concentration
in
XIX
6
cut
±
the
curvature
8
10“
circle:
g/ml
1
± 2
test,
g/ml
right:
G.A.;
in
the
triangle:
G.A.
into sections in orange
orange
1
light,
were
light.
length reached by the sections during the three
on
the
experiments
surface
were
of
the
made.
solutions,
Experiment
is
II
presented
is
plotted
in
in
THE
INFLUENCE
OF
RED
AND
FAR
RED
TABLE
Final
I.A.A.
in
acid
g/ml
in
0.
10
in
scale
27
SEEDLING
units
II
EXP.
Concentration of gibberellic
in
acid
g/ml
g/ml
0.
io-»
io-«
52.5
± 0.2
52.7 ±
0.1
53.1 ± 0.3
53.0
± 0.2
52.9 ±
0.3
53.8 ±
52.3
± 0.3
53.0
± 0.3
51.3
± 0.2
10
3.10“
sections
gibberellic
0.
1.10
AVENA
I
of
Concentration
concentration
THE
xx
length of
EXP.
ON
LIGHT
53.4 ± 0.2
0.3
53.3
53.4 ± 0.2
0.2
±
54.7 ± 0.2
52.8
±
0.2
53.3 ±
0.1
54.0
± 0.2
,J
53.7 ± 0.3
53.1
±
0.3
52.8
± 0.2
53.7
±
0.3
1.1th
8
54.1
0.3
53.5
± 0.3
55.3
±
0.3
54.1
±
0.2
3.10”
8
56.3 ± 0.6
55.5
±
57.4 ±
0.5
57.5 ± 0.5
i.io-
7
59.6 ±
0.7
60.3
± 0.7
I.10-
8
60.5 ±
0.7
60.2
±
1.10-°
3.10
Gibberellic
low I.A.A.
acid
±
indeed
enhances
and
concentrations,
0.4
the
of
growth
consequently
the
produces
0.8
sections
at
additional
an
peak in the graph.
Summary
The
test
of gibberellic
activity
well
as
as
its place in
however, is
This,
acid
the
sufficient
not
in
the
curvature
chromatogram
to
prove
are
the
CHAPTER
THE
IN
R.L.F.
shown
As
coleoptiles
latter
the
of
by
is
is
supplied
positive
be attributed
a
function
a
an
c.
an
are
THE
PHOTOTROPIC
phototropic
of the
short
light
time.
first
a
curvature
With
increase
increasing
positive,
in
negative and
a
response of cells
to
a
lateral shift of the auxin
synthesis
differences in
Now it
with
a
bearing
phototropic
at
higher light
intensities,
(1956).
on
the
behaviour,
By
none
of the
question of the
the
influence
light.
reviewed
were
investigators
cause
of red
of these
light
was
account.
was
shown in the
which
decreasing
I.A.A.
transport
auxin under the influence of the
to
and Went
experiments had
coleoptile
a
may
by the light,
the auxin
by Schrank (1950)
taken into
that
amounts
produced. These different responses
Arguments for and against each of these possibilities
whose
of Avcna
provided
energy
side,
inactivation of auxin
reduced
VIII
to
the shade
h.
a
curvature
redistribution of auxin due
to
d.
the
(1915)
within
of the R.L.F.
compounds.
two
AVENA COLEOPTIEES
light under these circumstances
second
a.
Arisz
periodic
a
OF
to those
of the
identity
CONNECTION WITH
REACTIONS
and in the straight-growth
test
comparable
is
preceding chapters
irradiated
I.A.A.
concentrations
with
light
concentrations,
before
it
is
of
but
reduced
X
that the
=
growth of
660 m/i
increases
again
to
This
zero.
a
decreases
at
low
relation
28
G.
tively by
The
for
curve
chapter,
I.A.A. concentration
of the
one
has been based
this
and
growth
between
shown
curves
the
plants that
in
will
be shown
on
9.
Let it be
the
As
blue
irradiated
Tentative
I.A.A.
side
the
light than
9.
Fig.
decreases
auxin than
side
pre-illuminated
increasing
the
I.A.A.
the shade side.
rate
level
auxin
larger
Now with
contain
at
be
can
At the outset
of the
than
the
well
as
I.A.A.
of unilateral
that of the
as
(at low light
the
however,
growth
rate
This
growth
of the shade side
this
curve
could
supposition
turning
the
(first
rate
of the
are
side
light
positive
from
light
the
once
may
curvature).
side
(negative curvature)
the tables
9
to
be
less
When
right side of the
may
more
the
surpass
whereas,
when the
turned
(second
curvature).
pre-illumination
a
growth
of
rate
turning
which
centration,
the
a
point
presents
the
by
same
the
checked
by investigating the influence
660
exercises on the
light of X
=
whose
negative
really
the
corresponds
rate
of
second
with
coleoptiles
irradiation with
series
to
phototropic
the relation between
growth
an
be
with
coleoptile
from
point
enhanced
In
the
peak has been passed,
which
growth
of
high but decreasing auxin
at a
of
slope towards the first peak
graph,
positive
energy, i.e.
the
coleoptile)
the
move
growth of the shade side
mounting
first
to
of
less
different
dot:
plants;
amounts
side
light
will
light.
increasing
of the
content
red
with
light.
amounts
coleoptiles
of intact
non-irradiated
content
of blue
quantities
receives
curve
of
irradiated
plants
in
of the
explanation
the illuminated side
the growth
discussed
31.
p.
coleoptile
side,
(solid line),
by the experiments
Fig.
imagined
along
graph
left, that of the light side preceding that of the shade side.
content
If
of
the I.A.A.
energy
shade
the
of
concentrations, circle:
level
light
of the
shade
course
with
tenta-
line).
assumed, with Oppenoorth (1941), that the
coleoptile
the
represented
(broken
Fig. 9 seems to offer a new possibility for
negative and the second positive curvature.
of
is
curve
Fig.
not
were
the data obtained
on
as
BLAAUW—JANSEN
of
light
experiments
of X
reaction
lies
positive
the
first
peak
of
growth and I.A.A.
should
=
at
the
at
curvature.
this
the
con-
point
be
660 m/r.
the influence which
irradiation
THE
with
INFLUENCE
light of 1
of these
RED
AND
FAR
RED
660 m// and of X
=
exercises
lengths
wave
studied
was
OF
also
in
the
=
of
THE
ON
480
29
AVENA SEEDLING
and with combinations
straight growth of coleoptiles,
the
on
regions
LIGHT
the
first
and
positive
negative
curvature.
METHOD
The
coleoptiles
experiments
a
particle
coleoptile
on
of
at
vermiculite
a
performed
was
serve
as
mark.
1
light of
660 mfi,
X
in
again.
was
The
described in
as
fixed
chapter II
by
cm
of X
=
the
560 m/i;
lanoline
tip;
the
this
with
unilaterally
the
old,
the
on
operation
particle
photographed
were
irradiated
of
means
from
for
88 hours
were
to
was
by
of
use
light of
X
—
both.
or
the irradiation
in
1
coleoptiles
and
m/t
increase
measured in
the
mfi,
Two hours after
light
green
Next
480
=
was
distance of about
560
=
cultivated
were
phototropism. When the seedlings
length
of
the
coleoptiles
the
were
photographed
from
coleoptiles
tip
to
mark
mm.
EXPERIMENTS
The
results
The result
Fig.
10.
of
of
one
Increase
in
these
experiments
of them is
length
in
plotted
arc
in
2 hours of the
of
Fig.
light
coleoptiles that were one-sidedly illuminated
sities: the coleoptiles of the lower row previously
those
of the
upper
row
not
recorded
side
in
Table
XXL
10.
(L)
with
and of the shade
blue
light
irradiated
previously
with
side (D)
of various inten-
light
irradiated.
of 660
rn.fi,
30
BLAAUW—JANSEN
O.
TABLE
in
Growth
during 2
mm
No
not
pre-illuminated
ergs/cm
2
A
=
660
m/i
not pre-illuminated
pre-illuminated with
2
700
A
ergs/cm
660
—
m/i
480
pre-illuminated
not
A
=
660
m/i
pre-illuminated
Transition
region
not
0.49
0.06
(16)
0.04
(23)
0.21
±
0.04
(17)
0.66 ± 0.06
(17)
0.57
± 0.05
(20)
0.20
±
0.04
(7)
0.59
±
0.43
± 0.07
(12)
0.13
±0.05
(9)
0.71
±0.09
0.40
±
negative phototropic
not
2
A
A
=
700
=
ergs/cm
660
A
not
(20)
0.60
±0.16
0.43
± 0.07 (12)
0.45
±
0.55
±
0.50
±0.17
=
at
a
(9)
A
480
=
shade
m/i
side
(8)
0.36
± 0.08
(8)
to
(18)
second positive
(8)
0.45
± 0.08
(8)
(3)
0.30 ±0.17
(3)
480
0.06
0.56 ±
(12)
ergs/cm
2
A
side
light
m/i
curvature
phototropic
24000
light of
=
0.08
0.05
480
=
m/i
shade side
(14)
0.56
± 0.05 (14)
rn/i
(15)
0.72
± 0.06
(15)
0.72
± 0.06
0.53
± 0.12
(06)
0.53
± 0.12 (06)
0.77
±
0.04
(17)
0.77
±
0.78
±
0.04
(11)
0.78
± 0.04 (11)
0.97
± 0.08
(12)
0.97
± 0.08 (12)
(15)
m/i
0.04
(17)
660
m/i
that:
700
with
in
region
ergs/cm
2
X
=
660
m
alone
fi
inhibits
tire
coleoptiles;
light of
but
in the
is
d.
0.06
negative
X
480
=
—
coleoptiles
hances in addition the
c.
2
side
light
± 0.05
irradiated with X
only,
(7)
with
of the
unilateral
ergs/cm
0.57
0.58 ± 0.05
660
irradiation
growth
b.
m/i
pre-illuminated
It appears
a.
480
0.08
with
pre-illuminated
2
±
curvature
4800
light of
=
0.50 ±
pre-illuminated
ergs/cm
m/i
shade side
with
pre-illuminated
700
side
light
480
(16)
from
pre-illuminated
pre-illuminated
2
=
0.04
A
700 ergs/cm
A
±
No
not
2
with
pre-illuminated
2
ergs/cm
0.12
0.06
No
ergs/cm
J
curvature
(8)
±
A
700
20
m/i
0.55
of the
Region
not
phototropic
irradiations
with
pre-illuminatecl
700
different
light of
-
A
after
hours
first positive
of the
Region
XXI
of the
2
m/<
660 m/i,
inhibits
pre-irradiated
growth
negative
if
(20 ergs/cm )
rate
at
curvature
given
growth
with
X
the shade
the
=
at
to
coleoptiles
the
660
light side
m/u,
it
en-
side;
growth of the shade side
inhibited;
the
turning
previous
point from negative
illumination
with
light
to
second
of X
=
positive
curvature
660 m/t enhances
the
THE
growth
OF
INFLUENCE
of the
rate
coleoptiles
AND
FAR
RED
the
coleoptiles;
also
is
RED
growth
than
larger
LIGHT
that
ON
THE
of these
rate
of
31
AVENA SEEDLING
controls
pre-illuminated
in
complete
of
unilateral
kept
darkness.
DISCUSSION
Table
blue
XXI
light
positive
This
light.
fits
introduction
in
they
9.
Fig.
the
the shade
at
according
schematically
irradiation
second
that
represented
light of X
the
transition
positive
the
growth
the
of
enhances
between
of
growth
and shade side of
the
light
660 m/./.
coleoptiles not irradiated with light of X
along the solid graph of Fig. 9, whereas the growth of coleoptiles
runs
shown
pattern
is
quantities
chapter
assumed that
I.A.A.
positive
auxin
blue
by
Terpstra
is
extraction
overall
that
by
a
irradiated with
of the
negative
The
positive
with red
lost
the
So
24 000
ergs/cm
2
X
one
than
the
to
this
=
itself
coleoptiles
is shown
a
which
character
since
estimated
that
after
thus
of
the
usual
chromatoshould be
far,
content
that is
was
coleoptiles
the transition
at
curvature.
XXI show that
curvature
of
method of auxin
(i.e.
advanced
480 m//,
negative
the inactivation of
manner
I.A.A.
the
the aid
by
prove
Oppenoorth
content
positive
right,
convincing
in
of the
light, nevertheless show
pronounced
growth
increasing
activity of the R.L.F.
Oppenoorth’s
evidence,
of Table
negative
their
that
I.A.A.
the
to
could
extracts
the
the second
to
the
of
the introduction
experiments
Secondly,
of his
determination
experiments
and
have
not
separation).
completed
the
be due
that he
reliable.
time, and
graphic
of
thought
content
in
decreases. If this is
must
demonstrated
not
auxin
influence
by them under the influence of
However,
coleoptile
light,
(1953)
by the broken
between
by increasing the quantities of blue light the
results
(1941)
under
that shown
curvature
the
Unfortunately,
Oppenoorth
indicated
resemblance
coleoptiles
of I.A.A.
of the
content
and second
660 m/t is
=
striking
and
light
decreasing
we
of X
a
the
by
of blue
quantities
light
there
Indeed,
line.
=
with
pre-irradiated
at
red
in
by the graph
region
So
curvature.
with
660 m/i
=
too
coleoptiles,
the
which
to
(first
side
expounded
was
with
in
for
true
pretreated
not
or
minimum
a
passes
growth
hypothesis
and
be
can
Moreover
and
negative
the
quantities
first
pretreated
are
the
growth of the coleoptiles
the
side
light
chapter
light and shade side
of
increasing
that then
with
this
to
the
(negative curvature). This holds
whether
matter
no
at
and
minimum
with
that
growth
curvature)
a
passes
shows
the
first
at
the
which
turning point of
growth peak,
by coleoptiles
irradiated
not
are
though
a
less
that have been irradi-
ated with
red light. This is indicated
by the solid line of Fig. 9. Presumably either decapitation and sectioning of the coleoptiles obliterate
this
less
same
From
of
pronounced
effect
the
Beyer
as
these
first
red
peak, or large quantities of blue light
light in producing a first growth peak.
experiments information is obtained also
positive
(1927)
the irradiated
that
curvature.
with
side of the
the
first
They
confirm
positive
coleoptile
is
the
curvature
inhibited,
the
on
have
the
cause
observation
the
growth
growth
the
at
of
at
the
32
G.
dark
side
this
the
on
other
enhancement
The
coleoptiles.
of
at
would
not
the
shade
red
light, is unlikely.
non-irradiated
I.A.A.
level
an
is
content
the
at
with
This
power
to
of
accumulation
end with
may
to
Arisz
coleoptile
With
reacts
positive
the
so
on
reacts
I.A.A.
of the
whether
called
I.A.A.
that
so
result
cannot
the
in
shade
side;
reduction of the
a
coleoptile,
of the
R.L.F.,
This is
which
possibly
in line
on
In
i.e.
stated
with
the
the
under
of
phototonus
a
the
that the
illumination
changes in
which
changes
influence
the
of
duration and
determine
a
the
the
way
following one-sided light stimulus.
a
light the plant
our
or
reach
can
by
a
negative
a
in
stage
curvature.
the
faculty to produce
opinion pre-illumination reduces
If it is
coleoptile.
positive
a
of the
contemplation
unilateral illumination
a
content
determines
to
side of the
undergoes
(1914)
curvature returns.
the I.A.A.
optimum
towards
region
the
opinion the first positive
longer duration of the pre-illumination
a
accumulation
Possibly
side
curvature
“tonus”),
with white
By pre-illumination
which it
our
of I.A.A.
all-round
preceding
a
the
light.
I.A.A.
short
a
or
stimulus
a
stimulus.
intensity of
in which
this
above the
So in
rate.
the action
(“Stimmung”
react
preceding
viz.
one,
(1956).
chapter
“disposition”
classic
been irradiated with
not
the shade
at
and shade
light
into
is
coleoptiles
shift
a
red
by
content
growth
both
an
views of Went
its
coleoptiles
to
coleoptile
combined
that
pre-irradiated
Maybe the difference in the reaction of irradiated
I.A.A.
due
brings
a
coleoptiles
negative and the second positive
I.A.A.
That
that have
enhancement of the
the
the
is
however,
accumulation of the available
an
coleoptile.
of the non-irradiated
curvature
in
only
explanation
causes
of
appears,
occurs
coleoptiles is connected with the reduction of the
increase of the
an
side
the
of the
content
light
side
with
and
shade
plausible
most
occur
hand enhanced. It
the
that the unilateral blue
auxin
BLAAUW—JANSEN
a
true
negative
coleoptile
must
that the I.A.A. level
will
curvature
entirely
result,
depend
upon
level.
Summary
In
this
curvature
due
negative and
a
to
the
can
gets
a
shift
second
reduction of the
factor
evidence
chapter
is
not
I.A.A.
only
adduced
in
level
the
for
to
the
INFLUENCE
ON
In
several
OF
an
on
dark
the
as
side
that
the
of the
rate
but
FAR
the
first positive
coleoptile;
other hand would
a
result
of
phototropic
be
the
which
influence growth. Consequently the
growth
THE
instances
hypothesis
the
coleoptile
CHAPTER
THE
the
to
curvature
positive
the opportunity
influence
is
of heteroauxin
red
red
light
as
response
the
due
to
light
factor
well.
IX
RED
LIGHT
(of
λ
740
=
mµ)
AVENA COLEOPTILE
antagonistic
effect
of red and
far
red
light
has been found.
In
(red
the
preceding
light)
the Avena
has
chapters
two
coleoptile,
it
different
was
and
shown that
probably
light
of A
independent
=
660 m/u
effects
on
THE
INFLUENCE
the
firstly:
maximum
OF
LA.A.
AND
RED
the
by
RED
of the
content
reached
FAR
LIGHT
ON
coleoptile
first
THE
is
positive
AVENA
the
decreased;
phototropic
33
SEEDLING
higher
is
curvature
presumably connected with this decrease;
secondly:
is
low
at
induced,
coleoptiles
I.A.A.
which
the
at
seemed
of X
light
perhaps
or
'l ire
the
worth
while
the
growth
of
rate
whether
investigate
to
(far red
maximum
growth of the
and
the second
light)
antagonizes
irradiation
of these
one
with
effects
both.
even
methods used
preceding
second
a
in
curvature.
740 m/i
=
increase
an
turning point between the negative
positive phototropic
It
concentrations
causes
this end
to
far
chapters
influence of red
the
were
the
same
those described in
as
corresponding
experiments
the
on
light.
EXPERIMENTS
First
effect
Avena
a.
cm
2
X
=
They
6000
resulting
to
ergs/cm
2
X
curvatures
produce
either
740 m/i
=
20 ergs/cm
2
X
or
without
m/i
9.0 ±
ergs/cm
700
6000
ergs/cm
2
X
2
ergs/cm
660
—
X
=
2
X
ergs/cm
It
2
X
m,u
740
that
appears
reverse
the effect
it
contrary,
The
b.
has
of the
light of X
not
the
subjected
growth
phototropic
2
cm
X
=
to
at
EXP.
17.1
12.2 ±
1.3
(10)
20.7
IV
11
Ill
EXP.
0.9
(10)
±
0.9
(7)
1.8
(08)
14.0 ±
1.8
(5)
± 0.8
(11)
13.4 ± 0.8
(5)
±
EXP.
12.3 ±
1.4
(7)
16.2 ±
1.3
(6)
9.9
the
it
viz.
—
can
be
1.4
of X
=
light of X
a
EXP.
V
9.9 ± 0.9
15.4 ±
light
with
effect,
same
8.6
±
1.1
(8)
(5)
14.0
±
0.8
(8)
740 m/i
=
drawn
from
on
inhibits
side
after
an
the
not
the
curvature.
the
on
the absolute
growth
curvature.
that
Like
were
XXIII), and it increases
coleoptiles
unilateral
On
experiments
growth of coleoptiles
light (Table
of
does
660 m/i.
larger maximum
740 m/i exercises
VI
(7)
region of the first positive
shade
curvature
480 m/t.
with
12.6 ±
unilateral blue
the
or
pre-illumination with
(09)
conclusion
660 m/i,
—
660 m/<
unilateral illumination
a
I
illumination with
in
a
(14)
fi
light of X
coleoptiles
by
1.4
of illumination
the
same
influence which
m
—
the
0.8
EXP.
6000
X
Table
+
m/i
740
In
2
ergs/
20
curvature.
XXII
±
13.3
m/r
660
=
ergs/cm
both.
after
or
EXP.
700
positive
XXII
induced
degrees)
480
=
700
with
with
shown.
are
(in
curvature
maximum
a
with
TABLE
Phototropic
irradiated
unilaterally
were
in order
m/«
prc-illuminated
were
with
coleoptiles
480
that
are
irradiation
developing
with
20
a
ergs/
34
G.
BLAAUW—JANSEN
TABLE
Growth
of the
shade
and
light
side
No
A
scale
light of
480
after
units)
Light
unilateral irradiation
side
Shade side
m/i
0.58
±
0.08
(20)
0.13
± 0.04 (14)
0.62
±
0.05
(14)
0.46
±
0.08
(12)
0.07
± 0.03
(11)
0.66
±
0.07
(11)
0.57
±
0.05
(20)
0.20
±
(
0.59
±
0.08
(7)
0.46 ± 0.09
(10)
0.22 ± 0.06
0.93
± 0.09
(12)
pre-illumination
no
=
XXIII
(in
pre-illuminated with
6000
ergs/cm
2
A
m/u
pre-illumination
no
pre-illuminated
6000
ergs/cm
2
I.A.A.
=
660
A
=
740
it
m/x
could
this
does;
that
proved
of Avena
content
m/i
be
is
shown
in
Non-irradiated
from
in
test
of
extracts
Coleoptiles
coleoptiles
irradiated with
6000 ergs/cm
2
7
=
tips
10
± 3
4
± 0.5
tips
tips
8
± 2
3
±
1
7 ± 2
1
±
1
studied
I.A.A.
the influence which
of the
peak
sections
660
=
at
m/x.
appears
that the
is
completely
light of
A
presents
Table
I.A.A.
that
XXV
neutralized
of
b.
=
ergs/cm
2
Finally
740 m/t
on
function
of
the
of the
irradiated
lengths
with
coleoptile
given in scale units.
are
of
exercises
light
of A
=
660 m/t
in solutions
9
a
of
I.A.A.
concentration of
3.10^
g/ml
9
g/ml
I
II
I
II
52.6 ± 0.2
51.2 ±0.2
51.4 ±0.3
51.3
0.2
52.7 ± 0.3
52.6 ±0.1
53.5 ± 0.3
52.8 ± 0.3
51.9 ±0.2
52.0 ± 0.3
52.3 ±0.1
±
0.3
with
A
=
660
m
53.3 ±
/<
with
pre-illuminated
ergs/cm
a
XXV
coleoptile cylinders
pre-illuminated
pre-illuminated
700
fi
been
final
effect
in
6000
have
the
m
as
by illumination of the seedlings
10~
ergs/cm
(compare chapter IV)
740
concentrations
TABLE
700
m/<
740 m/t.
=
Growth
2
test
=
growth
growth-enhancing
partly
or
light of A
coleoptiles
In
the critical
It
with
which
curve
concentration in
of A
light
740
effect
By application of the straight growth
a.
first
light of
as
way
30
Second
A
same
reduces
XXIV.
20
20
not
the
Table
740 m/t
=
XXIV
TABLE
we
7)
(12)
of A
light
coleoptiles
Curvature
Extract
0.04
with
Finally
c.
the
A
740
=
2
A
A
=
=
it
740
660
was
reverses
m
fi
m/i
±
investigated
the effect
whether
which
light
irradiation
of A
=
with
660 m/t
light
of
exercises
THE
INFLUENCE
of
growth
the
on
OF
RED
AND
FAR
RED
in
coleoptiles
the
negative and the second positive
coleoptiles
with
or
ergs/cm
ergs/cm
2
2
It
hilated
ergs/cm
2
X
660 m/r
=
=
740 m/t
and
then
X
=
480 m/i
(see
chapter
of the
length
be
not
can
irradiated
coleoptiles
TABLE
ergs/cm
6000
2
A
660
660
=
ergs/cm
2
A
24000
the
gives
this
treatment.
anni-
A
m/x
0.06
±
24000 ergs/cm
(11)
m/u
480
=
2
m/i
0.40
±
0.05
(15)
0.64
± 0.07
(15)
0.47 ± 0.07
(12)
+
m/r
740
=
480
with
of
light
=
0.45
700
with
660 m/{ is
=
irradiation
an
No
none
=
6000
by
XXVI
A
A
X
660 m/i
=
XXVI
after
light of
of
X
the
this end
followed
Table
hours
two
after
coleoptiles
of
Pre-illumination
2
To
740 m/<.
Growth
ergs/cm
between
region
curvature.
one-sidedly
VIII).
35
AVENA SEEDLING
immediately
doubted that the effect
by light of
700
transition
phototropic
X
in
increase
THE
ON
2
pre-illuminated either with 700 ergs/cm
were
700
LIGHT
m/r
DISCUSSION
Irradiation with
of
X
660 m/i
=
that
in
appears
I.A.A.
the
the
positive
found
X
740 m/i and with
effect
on
at
the
light,
blue
I.A.A.
that
low
as
they
It
content.
relation
a
growth rate,
I.A.A.
first
his
final
experiments
reversed
Now
red
compared
as
light,
function
of
of
3
Fig.
first
a
the
troduction of
Fig.
to
a
I.A.A.
were
of
a
similar
depression
of R.L.F. As irradiation
at
negative
antagonized
in favour
and second
negative
with
light of
exposed
not
curvature
that
the effects
to
and
red
by red
the
by
De Lint
irradiation
(1957).
somewhat
more
or
the
the
but it
by
irradiation
presents
accompanied
with
growth
by
the
as
a
in-
higher I.A.A. concentrations.
is caused
by the R.L.F.
pronounced the higher the
with far red
partly
yellow light.
which
is
on
decreased
coleoptiles,
introduction,
curve
viz.
light,
decrease of the
in probability.
content, gains
found also
far
the
on
hypothesis
the
of red
the
same
unilateral
which irradiation excercises
depression
is the
660 m/<
—
growth peak, i.e. the
curvature,
was
that the
in
X
is
too
concluded
concentration,
depression
5 shows that
and that the
some
induced
peak
of
however, have the
that of non-irradiated
shows
function of the
a
Illumination
positive
positive
coleoptiles,
the inhibition
our
first
the effect
on
of Avena
length
length
that
light
first
a
660 m/i,
and the decrease of the I. A. A.
A twofold effect of far red
In
—
the
between
of the
of
light
concentrations
between the
of the
concentrations.
on
as
light
region
phenomena
therefore may be
exists
growth
curvature
occurrence
light of X
have
the effect of
I.A.A.
finding is another argument
growth of coleoptiles,
enhancement
the
This
low
which
phototropic
with the
curvature
one
=
influence
link up the
attempt to
our
antagonizes
at
in the transition
740 m/i.
—
peak
which presents
The
positive
second
740 m/i
—
growth
curve
coleoptiles
on
by light of X
of
the
concentration.
exercises
and
light of X
on
extract
amount
light annihilates the first growth
36
G.
BLAAUW—JANSEN
the
peak caused by the R.L.F., it will, in all probability, also annihilate
growth depression due
the I.A.A. content of his
light
the
to
discussed
level
far
of the
content
red
far
a
red
coleoptiles
by irradiation with red
the
he
R.L.F.
has
light,
well
De
irradiation.
the
red
as
found
yellow
or
depression
with
reduces
far
the
that
do
they
red
I.A.A.
that the final
that have been
by coleoptiles
Lint
growth
depression
light,
impossible
be
must
attained
the
causes
abolished
as
it
coleoptile,
of the dark controls is
to
Presumably De Lint has reduced
where
and
above,
Since
light.
the R.L.F.
to
length
exposed
do this.
not
Summary
One
the
of
red
far
effects
two
growth
as
function
a
growth
of
rate
light
the
on
I.A.A.
red
into
has
light
two-peaked
a
the
WHICH
EXERCISED
the
AVENA
preceding
the
influence
is
further
order
DISCOVERY
RED
COLEOPTILES
it
chapters
reactions
exercised
of
light
of the
ORANGE
HAS
FOR
AUXIN
coleoptile
the
on
first
THE
light
and
I.A.A.,
leaf.
Without
performed
we
learn the effect of orange
THE
LIGHT
TESTS
on
primary
analysis
OF
AND
shown that orange and red
was
the
via
comprehensive
to
red
as
maximum
the
on
X
THE
BY
APPLICATION OF
In
one.
curvature.
CONSEQUENCES
influence
neutralized by
which presents
depressing effect
same
CHAPTER
ON
is
curve
coleoptile
one-peaked
non-illuminated coleoptiles and
positive phototropic
INFLUENCE
Avena
transformation of the
of the
far
On the other hand
the
of red
the
light, namely
some
light
that
this
intending
experiments
a
in
the results of the curvature
on
test.
I
Usually
room
the
on
last
the
switches
one
with
seedlings
orange
above,
the
intact.
If it
seedling
using
At
orange
the
to
were
light.
were
only
test
We
The
tested,
were
should
one
this
curvature
twice
removed after
applied
first
an
the first
to
series
of 30 minutes between the
that of the
first
rack
carried
interval
at
out
of
a,
to
treatment
of series
b.
as
two
decapitation.
next
the
to
been
react
shown
practice
operate
the
same
influences
when
carefully
next
experiments.
coleoptiles
hours.
b.
The
rack
were
primary leaf
subsequent
There
of the first
that
time with orange
usual, i.e. the plants
The
series
time
primary leaf remains
the racks with
illuminated
test was
with
in
the dark-
consequence
has
as
commercial
investigated
all
the
as
In
more
because,
long
so
much
have
ones
that this
plants,
light when entering
curvature test.
beginning of the experiment
be
decapitated
was
the first
shown
light.
a
decapitated
than
of the
the curvatures
out
carry
reacts
be
can
to
be
to
light
the orange
on
the intention
was
treatments
an
interval
of series
a
and
THE
INFLUENCE
OF
RED
AND
FAR
RED
TABLE
of
Effect
light
orange
LIGHT
in
g/ml
series
-1
0.
Table XXVII
is
curvature
used
is
to
to
apt
the
1
7
±
1
12 ±
1
±2
0.75
10~ 7
16
±
1
13
±
1
20 ±
1
16 ±
1
14
1
7
1.00
10~
1.25
10~7
15
agar
the
of
the
This
in
dark
b
the maximum
error
the
method is
test
of orange
of
till
hours after the
two
If this
a.
use
source
the
±
In series
series
to
auxin,
results.
plants
test
plants
with I.A.A.
compared
as
± 2
of the
curvatures
blocks
quantities
pervert
-2 ±
1
11
decreased
by keeping
b
10“
gives the
compare
±0
test
degrees
series
a
9 ±
7
in
0.50
of the
application
7
io-
0.25
37
SEEDLING
AVENA
(curvature)
Avena
Curvature
concentration
I.A.A.
THE
XXVII
the standard
on
ON
light therefore
can
be
eliminated
of de-
moment
very
capitation.
II
With
standard
the
tion. After
the first
though Van
Now
we
it
concentrations
second
did
the
decapitation
larger
or
find
to
der
Wey
by
second
a
leaf is
usually
that
decapitaloosened,
not.
or
“increase
fact
that
is
obtained)
were
circumstance
Der Wey
did this
whether the
denoted the
the
decapitate
to
finding of Van
whether he
out
curvatures
the
to
usual
primary
indicate
not
point
our
is
the
auxin is enhanced
to
which Van
(by
it
on
decapitation
Wey
der
made
sensitivity”
auxin
sensitivity
that the
(1931)
test
is based
curvature
coleoptiles twice; this practice
due
in
low
at
to
the
removal of the
the
primary leaf was postponed.
Four series
of
a
turn
I.A.A.
for
Series
of
plants
decapitation
a
primary
was
pulled
Series
c
after
1 i
after
hours
Series d
out
twice
the second
like
c,
but
with
series
same
series
of dilutions
kept in darkness till its
an
interval
of
1| hours. The
The
once.
series
the
but
a,
the
decapitation.
primary leaf
was
not
decapitation.
decapitation,
make this
to
was
in the usual way after the first
decapitated
was
with the
plants
of
come.
treatened like
loosened until after
immediately
had
decapitated
was
leaf was
Series b
tested
were
solution. Each rack
but
primary leaf
the
agar
comparable
I.A.A.
was
to
was
blocks
scries
applied
pulled
were
out
applied
a.
immediately
after
decapitation.
The results
The
are
largest
recorded in Table XXVIII.
curvatures
at
low I.A.A.
concentrations
found in series a, but this series shows the best
curvature and
I.A.A.
concentration.
are
Obviously
a
second
der
Wey
came
to
this conclusion
to
be
decapitation
does not enhance the reaction to low concentrations of I.A. A.
Van
not
proportionality between
by comparing
a
Probably
series
d with
38
G.
BLAAUW—JANSEN
TABLE
I.A.A.
the
of
g/ml
0.75
x
10-
± 2
13
±2
!
±
9 ± 2
7 ±
15
±2
1
10
± 2
± 2
x
io-
7
12
± 2
15
±
1
15
± 2
14
X
10-
7
16
± 2
15 ±
1
17
± 2
12 ± 2
7
17
±2
13 ±
1
18 ±
1
14 ±
1
19
±2
20
± 2
18
±
1
18
1
17 ±2
18
± 2
17
±
1
X
10-
x
10- 7
2.50
x
10-
b.
7
Indeed
of
amounts
a
series
responds
than series
vigorously
more
was
and
left
intact
not
by
the second
between series
a
decreases
decapitation
1| hours after
during
the
the
series
which
c,
to
b
beginning
appears
shows
instead
“sensivity”
d
in series
This
decapitation.
and
±
23 ±5
I.A.A., but this is caused by the fact that
comparison
second
b
j
1.50
2.00
!
1.00
illumination,
the
± 2
10
d
c
1.25
primary leaf
from
11
7
7
degrees
'
2
,
X
10- 7
in
1
b
a
0.50
series
small
Curvature
concentration
in
a
XXVIII
of
that
en-
hancing it.
So
if
plants
1
between
good proportionality
a
centration is
wanted,
needed,
are
should
one
should
one
and
curvature
I.A.A.
con-
decapitate twice, but if very sensitive
decapitate
and
once
apply
I.A.A.
\ hours after decapitation.
III
It
needs
apparently
no
illumination of the
test
likewise
of the
the results
special
plants with
experiments
orange
straight-growth
ascertain
to
red
or
that
light will influence
(cylinder)
test.
IV
It seemed worth while
influence
of
irradiation
positive
that the influence of
II
chapter
showed
we
maximum
the
enhances
question indicated above,
unilaterally with 20 ergs A
with
long after the illumination the
In
persists.
orange light
with
pre-illumination
a
appears
know how
to
light
curvature. To solve ihe
coleoptiles
after
orange
480
=
light.
orange
of
that
first
irradiated
we
1, 2, 4
0,
From
the
or
table
6 hours
XXIX
is still recognizable
orange light
after its application.
TABI.E
XXIX
Phototropic
Interval between
orange
light
irradiation with
blue
curvature
ergs/cm
2
1
in
=
degrees after
480
m/r
and
light
pre-illuminated
not
coleoptiles
pre-illuminatcd
coleoptiles
hour
1
(7)
15
±
1
(5)
1
hour
2H± 3
(5)
12
±
2
(7)
2
hours
1914- 2
(7)
14 ±
1
(7)
4
hours
19J±
2
(5)
1,3
G
hours
16
0
(5)
32 ±
±
!
0
I
with
20
pre-illumination
± 2
12 ±
it
six hours
1
(4)
(5)
THE
INFLUENCE
RED
OF
AND
RED
FAR
LIGHT
ON
THE
AVENA
dark
rooms,
39
SEEDLING
SUMMARY
1.
Orange
reactions
2.
the
The
Light
4.
Irradiation
it
a.
b.
reduces
6.
of A
the
R.L.F.
the
way
It
660
An
but
directly,
coleoptile
=
740
the
through
to be
appeared
m/t
2
whole, with 700 ergs/cm
a
but
m,«,
on
factor
A
active.
660
=
m/i
The
results
and red light
on
of
the
growth
which
ergs/cm
like
2
A
the latter,
and
experiments
our
m
the
of
are
as
effect
4b
of the
content
of
the
could
I.A.A.
with
that the influence of
should
not be
light
of
coleoptile.
curvature
positive
of the R.L.F.
tests
of
range
concentrations.
function
a
annihilates
ju
I.A.A.
warning
a
narrow
one.
the second
results of growth-substances
a
I.A.A.
growth
two-peaked
740
=
low
at
presents
a
in
coleoptile
rate
of the interaction
study
unidentified substance,
an
(R.L.F.).
the
curve
6000
coleoptiles.
the effect of
or
growth of the
enhances
reduces,
further
a
of the
content
explanation of the negative
be based
8.
as
transformed into
is
Irradiation with
7.
of A
light
as
amount
light
one-peaked
concentration
=
I.A.A.
regulates the
concentrations.
this
the
upon
influences
(I.A.A.).
primary leaf.
Avena seedling
called the red
The
I.A.A.
act
well
as
m/i
the
of
not
of the
tip
it enhances either the
here
5.
660
=
indole-3-acetic acid
effects:
two
I.A.A.
the
air-conditioned
the
to
does
light
of
of A
3.
in
used
as
coleoptiles
orange
intermediary
has
In
light,
of Avena
orange
underestimated
-
ACKNOWLEDGEMENT
Dr.
The author thanks Prof.
to
V.
investigation,
this
out
carry
for
J. Koningsberger
for
his
interest
in
providing
work
her
the
opportunity
his
and for
valuable
criticism.
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