The effect of environmental factors on growth. Development and alkaloid production of poppy (Papaver somniferum L.). I. Responses to day-length and light intensity


Bernath, J.; Tetenyi, P.

Biochemie und Physiologie der Pflanzen 174(5/6): 468-478

1979


Short-day rhythm hindered the stem growth (by 45-60%), the flower initiation (by 3-45 days), and the alkaloid accumulation (by 4-5 times in some cases) of Papaver somniferum in comparison with long-day program. Under long photoperiods increase of the light intensity (from 0.8 × 10^4 lux to 3.2 × 10^4 lux) accelerated growth and development of the three cultivars investigated: optimum growth occurs at IM 10^4 lux, whereas optimum development was observed at 3.2 × 10^4 lux. Similarly, the alkaloid biosynthesis increases 2-3 times at higher light intensity of long-day treatments (2,4-3.2 × 10^4 lux). At the same time the proportion of the higher methylated alkaloid components rises. In the ease of certain cultivars, depending on light condition, the presence either of morphine or codeine became characteristic. This change necessitates a new physiological aspect in intraspecific chemotaxonomic evaluation.

Biochem.
Physic!).
Pflanzen
174,
468-478
(1979)
The
Effect
of
Environmental
Factors
on
Growth.
Development
and
Alkaloid
Production
of
Poppy
(Papaver
somniferum
L.)
I.
Responses
to
Day
-length
and
Light
Intensity
JENO
BERN/Cali
and
PETER
TATENYI
Research
Institute
for
Medicinal
Plants,
Builakaitisz,
Hungary
Key
Term
Index:
morphinane-,
phtalideisoquinoline-,
benzyliso(plinolino
alkaloids,
alkaloid
biosynthesis,
effect
of
light;
Papaw
somilifcru»,.
Summary
Short
-day
rhythm
hindered
the
stem
growth
(by
45-60%),
the
flower
initiation
(by
3-45
days),
and
the
alkaloid
accumulation
(by
4-5
times
in
some
cases)
of
Papaw
soornifertem
in
comparison
with
long-day
program.
Under
long
photoperiods
increase
of
the
light
intensity
(from
0.8
10
4
lax
to
3.2
10'
hix)
accelerated
growth
and
development
of
the
three
cultivars
investigated:
optimum
growth
occurs
at
IM
10'
lux,
whereas
optimum
development
was
observed
at
3.2
10'
lux.
Similarly,
the
alkaloid
biosynthesis
increases
2-3
times
at
higher
light
intensity
of
long-day
treatments
(2,4-3.2
NI
lux).
At
the
same
time
the
proportion
of
the
higher
methylated
alkaloid
components
rises.
In
the
ease
of
certain
cultivars,
depending
on
light
condition,
the
presence
either
of
morphine
or
codeine
became
characteristic.
This
change
necessitates
a
new
physiological
aspect
in
intraspecific
chernotaxonomic
evaluation.
Introduction
New
results
were
achieved
in
the
biosynthesis
of
morphinane
alkaloids
(nrruEs
and
SCHi'TTE
1969)
as
well
as
by
practical
works
directed
towards
the
cultivation
of
Paper
rowel
-11e
species
(FENKA
1968.
i)Ir.MITRASKO
1977,
GILBERT
1977
etc.).
Importance
of
geographic
location
in
alkaloid
formation
by
LEE
CHANG
Kr
and
fivitNo
Hoot{
Kim
(1970),
the
weather
conditions
by
-
TETENVI
(1970),
TOROPOVA
and
LIMON
(1977),
Iforvx
and
NOVIKOVA(1968)
and
the
nature
of
soil
by
KUSEvIc
(1960)
have
been
report-
ed.
Physiological
effect
of
different
external
factors
(as
reflected
on
research
proposal
of
U.N.
Narcotics
Laboratory,
1975)
were
not
so
far
satisfactory
answered.
In
accordance
with
the
findiugs
of
TooKEv
et
al.
(1976)
we
intend
to
approach
the
changes
under
controlled
conditions
taking
into
consideration
the
interaction
of
light,
air
temperature.
nutrient
and
water
supply.
Our
aim
is
to
disclose
a
connection
between
growth,
development
and
alkaloid
formation
and
to
establish
their
relationship
as
well.
First
we
are
looking
for
what
extent
the
primary
metabolism
and
alkaloid
formation
can
be
effected
by
light
and
what
differences
may
be
found
in
the
behaviour
of
cultivars
differing
in
origin
or
in
alkaloid
chetnism.
Material
and
Methods
Final
ooaferial
The
investigations
were
carried
out
on
Hfingibrian
'K
6.k
Duna'
cultivar
and
thereupon
have
been
extended
to
cv.
`Kompolti
M
.
and
'Reading'.
Choice
of
the
cultivar
'Reading'
of
British
origin
Papaver
Alkaloid
Production
Affected
by
Light
469
was
justified
by
its
different
ecological
requirements
and
alkaloid
chemisin.
To
ensure
genetic
hoino-
genety
I,
seeds
were
used.
,ciandard
growth
conditions
Investigations
were
carried
out
in
phytotron
chambers
made
in
Canada
(E-15
and
modified
G-30
type
of
Conviron).
The
plants
were
grown
in
pots
on
per]ite
and
washed
soil
(v/v
1:
1).
After
the
germination
period
water
capacity
was
held
on
70%.
It
was
provided
by
watering
every
2
or
3
days
using
distilled
water.
During
germination
relative
humidity
of
the
air
was
adjusted
to
80%,
and
after-
wards
to
70
°„.
Nutrient
conditions
In
order
to
ensure
controlled
nutrient
supply,
plants
were
irrigated
weekly
with
KNOP
solution:
12.0
mg
N.
2.3
mg
P,
12.6
mg
K
(per
week
and
per
pot).
Dosages
were
doubled
from
the
beginning
of
the
differentiation
of
vegetative
apex
and
were
three
times
higher
from
beginning
of
the
budding
stage.
Temperature conditions
The
air
temperature
was
regulated
similarly
to
means
of
80
years
local
temperature
of
vegetation
period.
It
was
controlled
in
the
day/night
rhythm:
1st—
5th
week
12.5/7.3
CC
tith-11th
week
18.5/11.5
°C
12th
—17th
week
22.0/13.0
°C
18th--
26.0/16.0
°C
Light
coodilions
Levels
of
light
intensity
have
uniformly
been
set
by
General
Electric
F72T12
(CW)
V.H.0
fluores-
cent
lamps,
supplemented
by
incadescent
bulbs,
about
of
the
installed
wattage
calculated
on
energy.
Light
intensity
was
41)0
lux
during
germination.
Subsequently
light
treatment
variations
shown
in
rig.
1
have
been
applied.
Treatment
variations
have
been
chosen
to
achieve
the
maximum
production
achievable
at
the
given
intensity
level
anti
also
to
have
the
possibility
to
evaluate
some
of
the
lower
intensity
as
pre-trea
tinEuts
{light
conditions
prior
to
the
generative
differentiation
of
vege-
tative
apex).
10
h
illumination
cycle
was
applied
in
the
short
-day
and
14
h
in
the
long-day
treatments.
Number
of
treatment
variations
have
been
reduced
at
the
cv.
Kempolti
M'
and
'Reading.
Analysis
of
25
to
50
individuals
was
accomplished
at
all
treatments
and
all
eultivars.
Morpho-phenological
investigatinets
Growth
and
development
of
plants
was
evaluated
by
continuous
morpho-plienologieal
surveing.
Clime
of
germination,
vegetative
apex
differentiation,
bud
formation,
flowering
and
capsule
develop
-
0
B
Xl
IT
4
14x
1.6
x
70
4
/ur
2.4
x
1
3.2
x
ieridt
BEFe
R
E
GENctutrevE
A
F
Frl?
'ir
NreArroN
or
APEA
Fig.
1.
Light combinations applied in
the
course
of
the
experiments,
470
J.
BERNATH
and
P.
Tg:TP.Nvi
meat
was
established.
The
growth
was
characterised
by
continuous
measurement
of
height,
number
of
leaves,
flowers
anti
capsules.)
Final
processing
was
carried
oat
after
full
ripening.
Leaf
samples
were
taken
regularly
during
vegetation
period
to
follow
the
changes
of
alkaloid
composition
as
well
as
the
alkaloid
content
of
dry
capsule
was
analysed.
Alkaloid
analysis
The
qualitative
and
quantitative
determination
of
alkaloids
were
performed
by
double
-run,
thin
-
layer
chromatographic
method
of
Dittos
(1968):
0.2
g
of
capsules
dried
on
100
°C
and
powdered
were
measured
into
funnels
made
of
filter
paper.
Every
sample
was
percolated
ten
minutes
with
1
ml
sodiumcarbonate,
then
extracted
30
min
in
a
modified
Soxhlet
apparatus
with
20
ml
dinar
-
form
methanol
9:
1
(v/v].
Extracts
were
dried
and
residues
were
dissolved
in
1
nil
chloroform
methanol
1:
1
(v/v).
The
double
running
system
was
as
follows:
I.
benzene
methanol
9:
1
(viv
II.
chloroform
methanol
acetone
ethanol
6:
1:
I:
1
(vfv).
The
adsorbent
was
an
alkaline
Kieselgel
G
0.23
min
activated
at
110
°C.
Detection
was
carried
out
with
Dragendorff
reagent,
Evaluation
was
done
in
comparison
with
standards
of
known
quantities.
Three
repetitions
of
different
concentrations
of
every
sample
were
made
on
the
same
plate.
Results
Changes
of
growth,
development
and
dry
-mailer
production
Growth
of
the
plants
shows
an
characteristic
sigmoid
curve
(Fig.
2).
The
role
of
light
conditions
is
demonstrated
by
changes
of
ev.
'Kelt
Duna'.
The
period
of
internode
growth
extends
when
light
intensity
of
0.8
10
4
lux
(in
both
short
and
long-day
condi-
(cm)
100
50
r r
r r
1
70
id°
130
70
13.2
x
I
t"
/
t
,..
-
o
1
I
-
X.
.
-
t
I
A.'
A
/
ee
..,'
...-•
,•-•
do
s
100
130
70
100
116
x
et.
130
70
190
130
0.8
x
0.8
x
0'
!shrill
clJy1
Fig.
2.
Effect
of
Ugh!
conditions
on
Me
growth
measuring
by
the
height
of
vegetative
nr
generally
apex
location.
1.
pretreatment
of
0.4
10
4
lux
2.
pretreatment
of
0.8.
10
4
lux
(short
day)
3.
pretreatment
of
0.8.
10
4
lux
4.
pretreatment
of
1.G
10
4
lux
Papaver
Alkaloid
Production
Affected
by
Light
471
tions)
is
employed.
The
leight
intensities
of
1.6
10
4
and
3.2
10
4
lux
are
representing
the
optimum
light
conditions.
Also
the
effect
of
preatreatments
(No.
1-4)
can
be
demon-
strated
and
first
of
all
the
very
low
(0.4
.
10
4
lux)
intensity
and
the
short
-day
treatment
had
shortened
the
stem.
In
the
ease
of
`Kompolti
M'
and.
'Reading'
the
same
reaction
type
was
established.
Development
of
the
plants
was
essentially
determined
by
light
conditions
(Fig.
3)
also.
The
fastest
development.
occurs
at
3.2
10
4
lux.
Here
short
-day
pretreatment
(so.
2)
is
still
well
compensated
by
proper
postgrowing
conditions.
However,
on
plants
growed
at
1.6.
10
4
lux
and
usually
showing
a
slower
development,
a
short
day
and
very
low
intensity
pretreatment
(No.
1)
seems
to
be
critical.
Treatments
of
0.8
-
10
4
lux
affected
flowering
in
all
variation
unfavourably.
A
similar
tendency
could
have
been
observed
in
the
case
of
all
cultivars
investigated.
Differences
in
growth
and
development
were
unambiguously
shown
by
the
number
and
dimensions
of
capsules
provided
per
plant
(Table
1).
Also
in
this
regard
in
these
case
of
all
three
cultivars
the
critical
effect
occurs
at
1.6
10
4
lux.
Here
both
number
and
dimensions
of
the
capsules
decrease.
Their
calculated
"r"
value
is
0.859,
which
indicates
that
the
more
favourable
condition
reveals
itself
both
in
the
higher
number
and
size
of
capsules.
Differences
of
dry
-matter
production
also
depend
❑n
the
light
(Fig.
4).
The
smallest
differences
are
shown
by
the
roots.
The
production
and
proportion
of
stern
at
3.2
-
10
4
lux
is
high.
1.6
10
1
lux
marks
the
turning
point
where
the
rate
of
stem
and
leaf
equalizes.
Under
lower
illumination
and
short
-day
conditions
decreasing
the
total
production
the
proportion
of
leaf
becomes
predominant.
The
critical
quantitative
decrease
in
capsule
yield
takes
place
just
here
at
1.6
10
4
lux
(Fig.
5).
This
decline
becomes
general
at
0.8
10
4
lux.
The
highest
sensibility
is
shown
by
cv.
`Kompolti
M'.
+
3.2x1O
y
ivx
QA
CST
0.8
x
1
D
oule,„
shorl
day
Fig.
3.
Time
of
flowering
characterized
by
the
tiumber
of
days
elapsed
from
germination
1.
pretreatment
of
0.4
1W
lux
2.
pretreatment
of
0.8
10
4
lux
(short
day)
3.
pretreatment
of
0.8
10
4
lux
4.
pretreatment
of
1.6
1O
lix
31
Bio-oheni,
Physiol.
Pflanzem
Bd,
173
Table
1.
The
effect
of
day-lenght
and
light
intensity
cn
number
and
dimensions
of
capsulesfper
plant
Light
treatm
nt
(10
4
lux)
depending
of
the
time
of
generative
differentiation
of
apex
mean
number
of
capsules
(piece)
dimension
of
capsules
(cm)
primer
secondary
before
after
dimaeter
lenght
diameter
lenght
"KEK
DUNA"
"KOMPOLTI
M"
1.6
0.8
0.8
1.6
0.8
0.8
0.4
0.8
0.8
0.4
0.8
0.4
1.6
1.6
1.6
0.8
0.8
0.8
0.8
"READING"
1.6
1.6
1.6
0.8
0.8
0.8
0.8
short
day
short
day
short
day
short
day
short
day
short
day
short
day
short
day
3.2
2.44
3.3
2.7
2.3
1.9
2.55
3.6
2.6
2.1
1.'i
2.68
3.0
2.3
2.3
1.7
1.6
1.53
3.2
2.5
2.3
1.8
1.48
3.5
2.3
2.4
1.5
1.37
2.7
1.7
1.9
1.2
1.00
2.8
1.9
1.9
1.2
0.8
1.01
2.7
1.9
1.8
1.9
0.90
2
1
1.5
1.4
1.1
0.57
1.9
1.2
0.8
short
day
0.17
1.1
0.9
3.2
1.60
2.7
3.2
1.7
2.2
2.4
1.70
2.3
2.8
1.1
1.6
1.6
1.30
2.1
2.6
0.3
0.4
2.00
1.9
2.3
1.6
2.4
1.00
1.7
2.1
0.8
1.00
1.8
2.4
0.50
0.9
1.1
3.2
1.80
3.3
4.1
2.3
2.4
2.4
2.20
3.7
4.0
1.1
0.9
1.6
1.40
3.1
2.4
0.5
0.5
1.70
3.2
4.3
1.4
1.5
1.00
1.6
2.1
-
0.8
1.00
2.1
2.9
1.00
1.6
2.3
J.
BERNATH
and
P.
TgTENYI
Popover
Alkaloid
Production
Affected
by
Light
473
A
(g)
4,0
`i
3.0
0
2.0
1.0
1
.
12
8
x
/thud
19,8
X
lAirl
51
,
0f4
day
1
.6
x
?ehoj
nIt=i1=11
root
0
stern
0
leaf
Fir.
4.
Effect
of
light
conditions
(including
light
intensity,
day-lenght
and
their
conihinalions)
0/i
dry
wafter
prodoction
of
regelatire
organs
1.
pretreatment
of
(1.4
10
4
lax
2.
pretreatment
of
0.8
10
4
lux
(short
day)
3.
pretreatment
of
0.8
10
4
lux
4.
pretreatment
of
1.6
1(l
lax
).0
KEK
DUNA
1
short
day
1
0.8
2
4
1.6
(g)
KOMPOLT
1
M
'READING'
2D
a
gI°
EEO
3
PSULE
E
0
0
4
2
3
2
0
©
I
I:
T.EF
1.6
PT,
3,72
x
ter.
Fig.
5.
Effect
of
light
conditions
(including
light
intensity,
day
length
and
their
combikustionsfonidry
3
mailer
production
of
capsule
1.
pretreatment
of
0.4
11H
lux
2.
pretreatment
of
0.8
10
4
lux
(short
day)
3.
pretreatment
of
0.8.
10'
lux
4.
pretreatment
of
1.6
10'
lux
st*
4
474
J.
BERNATH
and
P.
TAT
ANY!
Quantitative
and
qualitative
changes
in
alkaloid
formation
At
leaf
rosette
stage
(with
10
to
11
leaves)
a
strong
accumulation
of
alkaloids
was
characteristic.
In
cv.
`Kompolti
W
and
`Kek
Duna'
the
presence
of
morphine
(80
to
85%
of
total
alkaloid),
while
in
cv.
'Reading'
the
presence
of
codeine
dominated.
After
generative
differentiation
the
total
alkaloid
formation
increased
changing
the
proportion
of
morphine.
Under
high
illumination
(2.4-3.2
10
4
lux)
the
presence
of
codeine,
while
under
low
light
intensity
the
presence
of
morphein
became
dominant.
The
amount
of
alkaloids
accumulated
in
dry
capsule
and
their
proportion
are
sub-
stantially
determined
by
the
ratio
changes
observed
during
growth
(Table
2).
Within
the
niorphinane
alkaloid
group
the
accumulation
becomes
more
intensive
with
the
increase
of
light
intensity
and
is
accompanied
by
a
characteristic
ratio
shift:
The
ratio
of
higher
methylated
components
(codeine,
thebaine)
increases.
This
is
well
demonstrated
by
change
of
accumulation
of
alkaloids
and
main
capsule
weight
pro-
jected
on
the
increase
of
light
intensity
(Fig.
6).
At
low
light
intensity
levels
resulting
low
capsule
weight
and
low
total
alkaloid
amount
still
the
morphine
provides
100%
of
inorphinanes,
thereafter
its
ratio
decreases
under
more
favourable
conditions
accom-
pa
ined
by
a
more
intensive
alkaloid
biosynthesis.
The
first
significant
accumulation
of
codeine
and
thebaine
occurs
at
1.6
10
4
lux,
but
more
significant
accumulation
can
be
seen
at
3.2
10
4
lux
only.
Whilst
the
increase
of
total
morphinalte
content
as
a
function
of
light
intensity
seems
to
be
a
general
tendency,
cultivar
differences
occur
in
the
ratio
change
of
compo-
nents
(Fig.
7).
Whereas
the
two
alkaloids
of
cv.
`Kompolti
M'
(morphine
and
codeine)
mg,
4
0
c
2
os
10
4
LUX
3.2
'KgK-DUNA`
M.
Co
a2
1
R
capsule
wt
(g)
Fig.
0.
S'imultaheous
changes
of
twilu
capsule
weight
and
alkaloid
acculnulation
processes
projected
on
the
increase
of
tight
intensity.
From
0.8.
10'
lax
to
3.2
10
4
lux.
M.,
morphine;
Co,
codeine;
Tb,
thebaine.
Table
2.
The
effect
of
day-lenght
and
light
intensity
on
qualitative
and
quantitative
composition
of
poppy
alkaloids.
Alkaloid
mgig
dry
matter.
Light
tr
atment
(10
4
lux)
depending
on
the
time
main
alkaloid
groups
of
generative
differentiation
of
apex
before
inorphinane
phthalide-,
benzyl-,
total
after
isoquinoline
morphine
codeine
thebaine
narcotine
laudanine
narcotoline
papaverine
`KEIC
DUNA'
1.6
0.8
0.8
1.6
0.8
0.8
0.4
0.8
0.8
0.4
0.8
0.4
`KOMPOLTI
M'
1.6
1.6
1.6
0.8
0.8
0.8
0.8
`READING'
1.6
1.6
1.6
0.8
0.8
0.8
0.8
short
day
short
day
short
day
short
clay
short
day
short
day
short
day
short
day
3.2
6.80
0.55
0.35
2.44
0.13
10.27
6.63
0.25
0.13
3.88
0.25
11.14
5.63
0.13
0.30
2.88
0.13
9.07
1.6
6.00
0.13
0.01
1.69
0.06
7.89
5.00
0.00
0.01
1.76
0.13
6.90
4.50
0.05
0.00
1.88
0.03
7.46
3.75
0.00
0.00
0.85
0.01
4.61
0.8
1.95
0.03
0.00
0.01
0.03
1.99
1.50
0.00
0.00
0.06
0.01
1.57
1.50
0.00
0.00
0.33
0.01
1.84
0.8
(short
day)
2.00
0.00
0.00
0.00
0.00
2.00
3.2
7.51
0.75
0.80
4.43
0.97
14.46
2.4
7.37
1.00
0.80
3.63
0.10
12.90
1.6
5.25
0.32
0.10
3.00
0.04
8.71
4.50
0.13
0.00
3.90
trace
8.53
2.75
0.08
0.00
3.25
trace
6.08
0.8
2.50
0.00
0.00
3.00
trace
5.50
3.2
3.50
5.68
0.25
2.81
5.15
17.39
2.4
2.80
5.00
0.33
3.05
5.05
16.23
1.6
2.75
4.60
0.02
3.08
4.33
14.78
2.90
3.33
0.05
2.75
4.00
13.03
1.25
0.13
0.00
0.12
1.75
3.25
0.8
2.50
0.28
0.00
0.62
3.63
7.03
3.07
0.83
0.00
0.50
3.10
7.50
Papaver
Alkaloid
Production
Affected
by
Light
14=.
Cn
476
J.
IIERNATa
and
P.
Ti:Tgxr:
show
the
sante
type
of
accumulation
as
described
at
cv.
'Kick
Duna',
'Reading'
gives
a
different
picture.
The
amount
of
morphine
is
growing
to
a
limit
value,
whilst
bio-
synthesis
and
accumulation
of
codeine
takes
place
continuously.
In
the
ease
of
plithidideisoquiooline
and
benzylisoquinoline
alkaloids
increase
of
total
amount
is
determined
by
light
conditions
can
be
revealed.
Yet
with
cv.
'KA
Dunit'
and
'Reading'
higher
variability
is
shown
by
phthalideisoquinoline
alkaloids
and
at
'hompolti
M'
by
benzvlisoquinoline
alkaloids.
Discussion
Poppy
alkaloids
can
not
be
regarded
as
curd
products
(Mu
LEI{
et
al.
1973;
FAIR
omits
et
al.
1978)
and
their
formation
and
accumulation
are
in
close
connection
with
primary
processes
(Buis
1978).
In
spite
of
that
fact
even
alkaloid
formation
was
investigated
mostly
with
full
disregard
of
plant
growth
and
development
processes.
Similarly
to
the
statement
of
GENTNER
et
al.
(1975)
we
proved
that
growth
and
development
of
plants
are
unfavourably
affected
by
short
-day
rhythm.
On
the
other
hand,
under
long-day
conditions
growth
and
development
are
clearly
accelerated
by
the
increase
of
light
intensity.
This
becomes
visible
in
the
increase
of
overal
production,
in
the
proportion
of
organs.
in
the
number
of
capsules
per
plant
and
in
the
size
and
weight
of
capsules,
too.
At
the
same
time
the
changes
are
on
a
small
scale
cultivar
dependent.
The
faster
development
and
increase
of
dry
-matter
production
affected
by
higher
light
intensity
is
accompanied
by
the
more
intensive
biosynthesis
of
alkaloids,
universally.
Under
that
conditions
probably
the
biosynthesis
of
higher
methylated
precursors
is
C
C
0
0
a
a
'KOMPOLTI
'
HO
0
6
4
2
Heo
a
HO
HP4e
'READING'
14e0
HO
f
r
1
TYPE
,'CHANGES
HO
NMe
NMe
03
t./3
02
capsule
wt.
(91
2.4
Fig.
7.
Ecological
oind
chemofoxonomical
differroce$
in
noorphioc-rodrior
sahiralion
CHM'
of
callicant
'EOM
POLTI
if'
and
'REAPING'
which
arc
raoscd
by
isioiallancoos
iriereakremcal
rrf
oupxote
weighl
mar
tight
inlenxily.
I,
morphine:
II,
codeine
Paparer
Alkaloid
Production
Affected
by
Light
477
promoted,
which
is
supported
by
the
fact,
that
more
significant
accumulation
of
reti-
culine
was
detected
at
2.4-3.2
10
4
lux
only.
At
higher
illumination
because
of
the
increase
of
precursors
and
methylation
processes
the
total
alkaloid
formation
rises,
which
can
be
detected
in
the
case
of
morphinane,
phthalideisoquinoline
and
benzyi-
isoquinoline
alkaloids
too.
Simultaneously
at
low
light
intensity
the
lower
alkaloid
content
seems
to
he
interpretable
by
absence
of
precursors
and
by
inhibition
of
methyla-
tion.
The
porportional
changes
of
components
can
be
considered
as
specific
process.
The
cause
of
it
can
be
searched
in
the
characteristic
demethylation
pathway
and
in
the
entalysing
enzyme
system
at
morphinanes.
At
low
illumination
the
precursors
of
morphine
group
are
formed
in
a
little
amount
and
are
able
to
demethylate
without
any
difficulties
almost
up
to
morphine.
On
the
contrary
the
larger
amount
of
precursors
can
be
demethylated
partially
when
the
overal
alkaloid
biosynthesis
is
promoted
by
light..
In
the
case
of
certain
cultivars
this
proportion
shift
results
differences
which
must
be
appreciated
from
chemotaxonomic
point
of
view.
All
three
investigated
culti-
vars
are
of
morphine
character
at
Iow
light
intensity,
whilst
"Reading"
cultivar
shows
an
increasing
codeine
character
due
to
its
low
morphine
saturation
level
when
light
intensity
becomes
higher.
This
also
refers
to
the
observations
of
TOMMY
et
al.
(1976)
who
could
not
interpret
the
increased
codeine
production
of
plants
raised
in
phytotron.
Although
we
have
supposed
the
importance
of
external
factors
on
the
for-
mation
of
alkaloids
earlier
(TfrdNyi
and
VAGITJFALVI
1965;
TETANYI
1970)
which
was
supported
by
other
theoretical
and
practical
observations
(PFETFR
1962;
HoTYN
and
NOV1KOVA
1968;
JVANOVA
and
GAEVSKIJ
1970,
Research
Recommendation
of
U.N.
Narcotics
Laboratory
1976
etc.)
it
could
be
revealed
exactly
for
the
first
time
by
us
how
the
growth,
development,
alkaloid
and
dry
matter
formation
and
accumulation
ogpoppy
"
11
1
)1PhYSIOlOgiCailY
affected
by
light.
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D.:
Biosynthesis
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B.:
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2011
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