Application of the polyamine putrescine increased yield of 'Mauritius' litchi (Litchi chinensis Sonn.)


Stern, R.A.; Gazit, S.

Journal of Horticultural Science and Biotechnology 75(5): 612-614

2000


During the last 5 years (1994-98, Israel), we have studied the effect of putrescine applied at the beginning of female bloom on litchi cv. Mauritius fruit yield. Required concentrations varied from year to year, but consistent increases were obtained with 5 x 10-5, 2.5 x 10-4 and 5 x 10-4 M putrescine. In 1994, inflorescences were immersed in a putrescine dichloride (putrescine) solution at 5 x 10-6, 5 x 10-5, 5 x 10-4, or 5 x 10-3 M. The 2 intermediate concentrations significantly increased the number of fruits per panicle, by about 100%. In 1995, individual trees were sprayed with putrescine at 1 x 10-5, 5 x 10-5, 2.5 x 10-4, 5 x 10-4, or 5 x 10-3 M. The best effect was obtained at the 3 intermediate concentrations. In 1996, individual trees were sprayed with putrescine at 5 x 10-5, 2.5 x 10-4, or 5 x 10-4 M (8, 40 or 80 ppm, respectively). A significant yield increase of about 50% was obtained with all treatments. In 1997 and 1998, large-scale trials were conducted, using 40 ppm putrescine. In 1997, yield was increased by 102%, from 4.1 to 8.3 tonnes/ha. In 1998, yield was increased by 60%, from 7.2 to 11.5 tonnes/ha. Weight of individual fruit was not significantly affected in any of the trials.

Journal
of
Horticultural
Science
&
Biotechnology
(2000)
75
(5)
612-614
Application
of
the
polyamine
putrescine
increased
yield
of
'Mauritius'
litchi
(Litchi
chinensis
Sonn.)
By
RAPHAEL
A.
STERN
1
*
and
SHMUEL
GAZIT
2
1
MIGAL,
Galilee
Technology
Center,
P.O.
Box
90000,
Rosh
Pina
12100,
Israel
2
The
Kennedy-Leigh
Centre
for
Horticultural
Science,
The
Hebrew
University
of
Jerusalem,
Faculty
of
Agriculture,
Rehovot
76100,
Israel
)
(Accepted
30
April
2000)
SUMMARY
During
the
last
six
years,
we
have
studied
the
effect
of
putrescine,
applied
at
the
beginning
of
female
bloom,
on
'Mauritius'
fruit
yield.
Concentrations
required
varied
from
year
to
year,
but
consistent
increases
were
obtained
with
5.10
-5
,
2.5.10
and
5.10
-4
M
putrescine.
In
1994,
inflorescences
were
immersed
in
a
putrescine
dichloride
(putrescine)
solution
at
5.10
-6
,
5.10
-5
,
5.10
-4
,
or
5.10
-3
M.
The
two
intermediate
concentrations
significantly
increased
the
number
of
fruits
per
panicle,
by
about
100%.
In
1995,
individual
trees
were
sprayed
with
putrescine
at
1.10
-5
,
5.10
-5
,
2.5.10,
5.10
-4
,
or
5.10
-3
M.
The
best
effect
was
obtained
at
the
three
intermediate
concentrations.
In
1996,
individual
trees
were
sprayed
with
putrescine
at
5.10
-5
,
2.5.10,
or
5.10
-4
M
(8,
40
or
80
ppm,
respectively).
A
significant
yield
increase,
of
about
50%,
was
obtained
with
all
treatments.
In
1997
and
1998,
large-scale
trials
were
conducted,
using
40
ppm
putrescine.
In
1997,
yield
was
increased
by
102%,
from
4.1
to
8.3
t
ha
-1-
.
In
1998,
yield
was
increased
by
60%,
from
7.2
to
11.5
t
ha
1
.
Weight
of
individual
fruit
was
not
significantly
affected
in
any
of
the
trials.
I
4
itchi
produces
a
large
excess
of
female
flowers
.
compared
with
the
number
needed
to
set
a
heavy
crop.
Nevertheless,
a
profuse
bloom
frequently
ends
in
a
poor
yield
(Menzel,
1984).
Massive
fruitlet
drop
usually
starts
after
the
end
of
the
female
bloom.
A
first
wave
of
flower
and
fruitlet
drop
begins
at
the
end
of
the
female
bloom
and
continues
for
about
a
month.
About
90%
of
all
fruitlets
abscise
during
this
period
(Stem
et
al.,
1995,
1997a).
Most
of
the
abscised
fruitlets
are
devoid
of
embryo
or
have
an
aborted
embryo
(Menzel,
1984).
A
second
fruitlet
drop
occurs
during
the
period
of
rapid
embryo
growth,
which
occurs
in
'Mauritius'
about
5-7
weeks
after
fruit
set.
Most
fruitlets
dropped
during
this
wave
have
an
embryo
(Stern
et
al.,
1995).
In
a
series
of
studies,
we
found
that
we
can
significantly
reduce
this
drop
by
spraying
with
synthetic
auxin
before
the
start
of
rapid
embryo
growth
(Stem
et
al.,
1995,
1997a;
Stem
and
Gazit,
1997).
In
'Mauritius',
this
treatment
usually
ensures
a
good
yield.
However,
sometimes
almost
all
the
fruitlets
drop
during
the
first
wave,
resulting
in
a
poor
yield
even
with
the
later
standard
auxin
spray.
The
underlying
reason
for
this
excessive
drop
may
be
climatic
(extreme
hot
or
cold
spells),
but
usually
it
seems
to
be
the
result
of
fertilization
failure
and
embryo
abortion
(Menzel,
1984).
Hence,
it
is
crucial
to
ensure
adequate
pollination
and
a
successful
fertilization
process.
Polyamines
have
been
found
to
increase
fruit
set
and
yield
in
several
fruit
crops
(Costa
and
Bagni,
1983;
Rugini
and
Mencuccini,
1985;
Crisoto
et
al.,
1988;
Singh
and
Singh,
1995),
including
litchi
(Mitra
and
Sanyal,
1990).
In
the
present
work
we
determined
the
effect
of
the
polyamine
putrescine,
applied
at
the
start
of
female
*Author
for
correspondence.
bloom,
on
'Mauritius'
yield.
The
recommended
auxin
spraying
was
performed
about
five
weeks
after
the
putrescine
treatment.
MATERIALS
AND
METHODS
The
trials
were
conducted
in
two
young
commercial
`Mauritius'
litchi
orchards.
Four
experiments
were
conducted
from
1994
to
1997
at
the
Lavi
orchard
(250
m
above
sea
level,
in
the
Galilee
mountains),
planted
in
1989
with
4
X
5
m
spacing,
and
one
experi-
ment
was
conducted
in
1998
at
Genosar
orchard
(100
m
below
seal
level,
in
the
Jordan
valley),
planted
in
1990
with
the
same
spacing.
The
orchards
were
"fertigated"
during
the
dry
season
(from
April
to
September)
and
water-stressed
in
October-November,
until
the
start
of
the
rainy
and
cool
winter
(Stem
et
al.,
1993).
The
trees
in
each
orchard
were
uniform
in
size,
healthy
and
flowered
profusely
in
April.
At
the
beginning
of
June,
when
the
fruitlets
weighed
about
2
g,
and
just
before
the
second
wave
of
abscission,
the
trees
were
sprayed
with
0.15%
Tipimon,
containing
100
ppm
of
the
synthetic
auxin
2,4,5-TP
(Stern
et
al.,
1995).
Preliminary
trials
In
1994,
litchi
inflorescences
on
five
year
old
trees
in
the
Lavi
orchard
were
immersed
for
10
s
in
51
plastic
buckets
containing
putrescine
(1,4-diaminobutane
di-
hydrochloride;
Sigma
Chemical
Co.,
St.
Louis)
solution
at
four
concentrations:
5.10
-6
,
5.1e,
5.10
-4
,
and
5.10
-3
M.
The
surfactant
Tween
20
was
added
to
all
solutions
at
0.1%.
All
the
treatments
were
applied
at
the
beginning
of
female
flowering.
The
experimental
design
consisted
R.
A.
STERN
and
S.
GAzn
613
TABLE
I
The
effect
of
immersing
'Mauritius'
litchi
inflorescences
at
the
beginning
of
female
bloom,
in
different
concentrations
of
putrescine,
on
number
of
fruits
per
panicle
and
fruit
weight
Data
are
the
means
of
10
inflorescences
(10
trees
X
one
inflorescence).
The
experiment
was
conducted
at
Lavi,
in
1994
Putrescine
concentration
Fruits/panicle
Fruit
wt
(g)
(M)
(ppm)
(no.)
0 0
9b
22.5
a
5
X
10
-6
0.8
11
b
23.0
a
5
X
10
-5
8
18
a
22.2
a
5
X
10
-4
80
20
a
22.4
a
5
X
10
-3
800
13
b
22.9
a
Results
within
a
column
followed
by
different
letters
differ
significantly
by
Duncan's
multiple
range
test,
P
=
0.05.
of
10
randomized
blocks;
one
tree
per
block,
one
inflorescence
per
treatment.
Trials
with
individual
trees
In
1995
and
1996,
six
and
seven
year
old
trees
in
the
Lavi
orchard
were
sprayed
using
a
motorized
back-
sprayer
(Echo
Air
Blower
Model
DM9),
21
per
tree.
The
experimental
design
consisted
of
five
randomized
blocks,
with
one
tree
per
block
in
1995,
and
ten
randomized
blocks
with
one
tree
per
block
in
1996.
At
harvest,
yield
was
determined
per
tree.
Average
fruit
weight
was
determined
by
weighing
50
randomly
sampled
fruits
per
tree.
Commercial-scale
trials
In
1997
(at
Lavi)
and
1998
(at
Genosar),
eight
year
old
trees
were
sprayed
with
a
commercial
1000
1
Spidet
blower
sprayer.
The
volume
of
spraying
solution
was
500
1
ha
-1
.
The
experimental
design
in
both
orchards
consisted
of
four
randomized
blocks,
with
50
experi-
mental
trees
per
block.
At
harvest,
yield
was
determined
per
tree.
Average
fruit
weight
was
determined
by
weighing
20
randomly
sampled
fruits
per
tree.
Statistical
analysis
Data
were
analyzed
for
statistical
significance,
using
the
general
linear
model
(GLM)
procedure.
Duncan's
multiple
range
test
was
used
to
compare
treatments
when
ANOVA
showed
significant
difference
among
the
means.
TABLE
II
Effect
of
five
concentrations
of
putrescine,
sprayed
at
the
beginning
of
female
bloom
with
a
motorized
back-sprayer,
on
yield
and
fruit
weight
of
`Mauritius'
litchi.
Data
are
the
means
of
five
replicate
trees
per
treatment
(five
blocks
X
one
tree
per
block).
The
experiment
was
conducted
at
Lavi,
in
1995
Putrescine
concentration
Yield
(kg/tree)
Fruit
wt.
(g)
(M)
(ppm)
0
1.0
x
10
-5
5.0
X
10
-5
2.5
X
10-4
5.0
X
10
-4
5.0
X
10
-3
0
1.6
8
40
80
800
5.2
a
4.9
a
8.6
a
7.1
a
6.4
a
3.4
a
243
a
23.9
a
24.1
a
243
a
27.6
a
24.1
a
Results
within
a
column
followed
by
different
letters
differ
significantly
by
Duncan's
multiple
range
test,
P
=
0.05.
RESULTS
Inflorescence
immersion
Putrescine
at
5.10
-5
and
5.10
-4
significantly
increased
by
about
100%,
the
number
of
fruits
per
panicle
(Table
I).
No
significant
increase
was
found
at
the
lowest
(5.10
-6
M)
and
highest
(5.10
-3
M)
concentrations.
Fruit
size
was
not
affected
by
any
treatment.
Trials
with
individual
trees
Yield
was
noticeably
higher
for
the
three
intermediate
concentrations,
whereas
it
was
lower
for
the
highest
and
lowest
ones
(Table
II).
However,
differences
were
not
significant
(P
=
0.07),
probably
because
of
the
small
number
of
trees
(five)
per
treatment.
In
1996,
we
used
only
the
three
putrescine
concentra-
tions
that
were
effective
in
1995.
In
addition,
we
determined
the
effect
of
three
surfactants
at
the
intermediate
concentration
of
2.5.10
M
(40
ppm).
Putrescine
at
5.10
-5
,
2.5.10
and
5.10
-4
M
caused
a
similar
significant
yield
increase
of
about
50%,
with
no
effect
on
fruit
size
(Table
III).
No
significant
difference
was
found
among
the
three
surfacants.
Commercial-scale
trials
In
1997
at
Lavi
orchard
and
in
1998
at
Genosar
orchard,
40
ppm
putrescine
with
the
surfactant
Triton
X-
100
(0.025%)
was
sprayed
commercially.
In
both
years,
the
putrescine
significantly
increased
'Mauritius'
litchi
yield
-
by
102%
and
60%,
respectively,
with
no
effect
on
fruit
size
(Table
IV).
TABLE
III
Effect
of
spraying
putrescine
at
three
concentrations
and
with
three
surfactants
at
2.5
10
-4
M
(40
ppm)
at
the
beginning
of
female
bloom,
with
a
motorized
back-sprayer,
on
yield
and
fruit
weight
of
'Mauritius'
litchi.
Data
are
the
means
of
10
replicate
trees
per
treatment
(10
blocks
X
one
tree
per
block).
The
experiment
was
conducted
at
Lavi,
in
1996
Putrescine
treatments
Surfactant
Yield
Fruit
wt.
(g)
(M)
(ppm)
(kg/tree)
Increase
(%)
0 0
12.7
b
24.1
a
5.0
X
10
-5
8
Tween
20
19.3
a
52
23.8
a
(0.1%)
2.5
X
10
-4
40
Tween
20
20.2
a
59
23.6
a
(0.1%)
2.5
X
10
-4
40
Triton
X-100
18.9
a
49
24.0
a
(0.025%)
2.5
X
10
-4
40
Shatah
90
21.0
a
65
24.3
a
(0.025%)
5.0
X
10
-4
80
Tween
20
18.5
a
46
23.8
a
(0.1%)
Results
within
a
column
followed
by
different
letters
differ
significantly
by
Duncan's
multiple
range
test,
P
=
0.05.
614
Putrescine
and
litchi
yield
TABLE
1V
Effect
of
commercial spraying
(with
a
1000-1
Spidet
blower
sprayer)
with
40
ppm
putrescine
on
yield
and
fruit
weight
of
8-year-old
trees
in
two
`Mauritius'
litchi
orchards.
Data
are
the
means
of
200
trees
(four
blocks
X
50
trees)
per
treatment
Yield
Treatment
(tonne/ha)
Increase
(%)
Fruit
wt.
(g)
Lavi
1997
Control
4.1
b
-
22.6
a
Putrescine
83
a
102
22.8
a
Genosar
1998
Control
7.2
b
23.0
a
Putrescine
11.5
a
60
23.3
a
Results
within
a
column
followed
by
different
letters
differ
significantly
by
Duncan's
multiple
range
test,
P
=
0.05.
DISCUSSION
Putrescine,
applied
to
inflorescences
and
whole
trees
at
the
beginning
of
female
bloom,
significantly
increased
`Mauritius'
yield
(Tables
I,
III,
IV).
Mitra
and
Sanyal
(1990)
found
a
similar
significant
increase
after
spraying
inflorescences
of
'Bombay'
twice,
"before
anthesis
and
into
open
flowers".
However,
they
found
that
fruit
size
and
weight
decrease
after
treatment
with
putrescine.
In
contrast,
in
our
trials
(Tables
I-IV),
no
effect
of
putrescine
on
fruit
weight
was
discerned.
The
different
responses
may
be
related
to
specific
cultivar
suscept-
ibility
and/or
to
the
extra
spraying
in
their
work
with
the
very
high
surfactant
concentrations
of
0.5
to
1%.
The
consistent
positive
effect
of
putrescine
found
in
our
trials
justifies
further
research
to
determine
the
optimal
putrescine
concentration
(apparently
between
8
and
40
ppm)
in
'Mauritius'
and
other
litchi
cultivars,
and
the
effects
of
other
polyamines
(Singh
and
Singh,
1995).
In
all
our
trials,
2,4,5-TP
was
sprayed
about
five
weeks
after
peak
female
bloom
to
reduce
the
second
wave
of
fruitlet
drop
(Stem
et
al.,
1995).
Thus,
we
can
tentatively
conclude
that
putrescine
improved
the
set
of
normal
fruitlets
and
may
also
have
decreased
fruitlet
drop
during
the
massive
drop
occurring
in
the
first
month
(Stem
et
al.,
1995).
The
positive
effect
of
putrescine
on
fruit
set
may
be
attributed
to
its
positive
effect
on
the
fertilization
process.
Polyamines
are
known
to
inhibit
ethylene
biosynthesis
(Apelbaum
et
al.,
1981;
Kushad
et
al.,
1988;
Tiburcio
et
al.,
1993)
and
in
this
way
retard
pistil
senescence
(Nichols
et
al.,
1983;
Faust
and
Wang,
1992;
Stead,
1992;
Lindstrom
et
al.,
1999).
The
litchi
pistil
retains
its
viability
for
about
5
d
(Stem
et
al.,
1997b).
`Mauritius'
is
the
first
cultivar
to
flower
in
the
commercial
orchards
of
Israel.
Hence,
it
depends
for
pollination
mainly
on
pollen
produced
by
M
1
and
M2
`Mauritius'
flowers
and
M
1
'Floridian'
flowers
(Stem
and
Gazit,
1996).
The
M
1
'Mauritius'
bloom
overlaps
some-
what
with
the
beginning
of
the
female
bloom.
At
that
stage,
female
flowers
are
not
fully
mature
(Stem
et
al.,
1997b)
and
in
addition,
the
viability
of
'Mauritius'
M
1
pollen
is
quite
low
(Stem
and
Gazit,
1998).
Thus,
effective
pollination
occurs
mainly
at
the
end
of
the
female
bloom.
Prolonging
pistil
viability
would
result
in
a
much
longer
effective
pollination
period.
We
suggest
that
this
is
the
main
mechanism
by
which
putrescine
improves
fruit
set.
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