Plant Physiol.
(1975) 55, 322-327
Abscission: The Initial Effect of Ethylene Is in the Leaf Blade'
Received for publication August 14, 1974 and
ELMO M. BEYER, JR.
Cenitral Research Depai-tinent, Experimental Stationi, E. I. dii Pon2t de Nenmouris anid Comiipaniy, Wilmington,
Delaware 19898
ABSTRACT
The leaf blade of cotton (Gossy-piu7n hirsutum L. cv.
Stoneville 213) was investigated as the initial site of ethylene
action in abscission. Ethylene applied at 14 ul/l to intact 3-
week-old plants caused abscission of the third true leaf within
3 days. However, keeping only the leaf blade of this leaf in
air during ethylene treatment of the rest of the plant com-
pletely prevented its abscission for up to 7 days. This in-
hibition of abscission was apparently the result of continued
auxin production in the blade since (a) the application of an
auxin transport inhibitor to the petiole of the air-treated leaf
blade restored ethylene sensitivity to the leaf in terms of
abscission; (b) repeated applications of naphthaleneacetic
acid to the leaf blade of the third true leaf, when the entire
plant was exposed to ethylene, had the same preventive effect
on abscission of this leaf as keeping its leaf blade in air; and
(c) the inhibitory effect of ethylene on auxin transport in the
petiole, which is reduced by auxin treatment, was also re-
duced by placing the leaf blade in air.
The reverse treatment of exposing only the leaf blade of the
third true leaf to 14 ,l/l of ethylene, while the rest of the
plant was kept in air, also did not cause abscission for up to
5 days. Auxin transport in the petioles of these leaves, how-
ever, was inhibited over 80% within 2 days and this effect
presumably accounted for their increased sensitivity to ethylene
during the subsequent exposures of the whole leaf to the gas.
These results suggest that an initial and essential function
of applied ethylene in abscission is to reduce the amount of
auxin transported out of the leaf blade. This reduction to-
gether with the inhibitory effect of ethylene on auxin trans-
port in the petiole reduces the auxin level at the abscission
zone to a point where the cells in this region become responsive
to the more direct action of the gas (e.g., enzyme induction
and secretion). This sequence of events accounts for the lack
of abscission unless ethylene is applied to both the leaf blade
and the abscission zone.
Auxin and ethylene are without doubt two of the more im-
portant regulatory agents involved in the natural control of
abscission (3, 25, 32). The dominant role of auxin in abscission
is one of retardation, while that of ethylene is clearly one of
promotion. The interplay between auxin and ethylene in their
mutually antagonistic roles in abscission has been the subject
of numerous studies. From these studies has emerged the
' Central Research Department Contribution No. 2174.
322
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in revised form October 4, 1974
concept that the ability of ethylene to initiate cell separation
in the abscission zone through enzyme induction (2, 4, 23, 34)
and secretion (5) depends largely upon the sensitivity of this
tissue to ethylene (1, 3, 16). The principal factor governing
this sensitivity to ethylene seems certainly to be auxin. Several
types of evidence point to this conclusion. First, auxin applied
to freshly cut petiole explants (16, 27). debladed petioles (3,
21), intact plants (9), or plants with their roots removed (21),
prevents ethylene induced abscission. Second, lowering the
auxin level at the abscission zone by excising and aging the
abscission zone tissue (1, 6, 16), removing the leaf blade (3),
or applying an inhibitor of auxin transport to the petiole (8,
28, 30) potentiates the action of ethylene.
If the auxin level at the abscission zone must first be lowered
before ethylene can induce cell separation, the question arises
as to the mechanism(s) and target tissue(s) involved in this auxin
lowering process when intact plants are exposed to the gas.
While several studies (9, 11, 15 18, 22, 36) have dealt with the
possible mechanisms involved in the reduction of auxin (e.g.,
transport inhibition, destruction, synthesis), none have specifi-
cally dealt with the problem of defining in the intact plant the
target tissue where these processes principally occur. The
technique of isolating and keeping one of the leaf blades of
intact cotton plants in air while treating the rest of the plant
with ethylene, or vice versa, was used in the present study to
answer this question.
The experiments reported here demonstrate that the leaf
blade is the initial target tissue of exogenously applied ethylene,
where some essential function of ethylene must first be per-
formed before abscission can occur. Results indicate that this
essential function of ethylene in the leaf blade is to reduce the
amount of auxin transported out of the blade. A preliminary
report of this study has been published (12).
MATERIALS AND METHODS
Plant Culture. Cotton (Gossvpiluml Ihirsutumt L. cv. Stone-
ville 213) plants were grown in a controlled environmental
growth room (2200 ft-c, 14-hr photoperiod, relative humidity
50 ± 5% day and night; temperature 27 C day, 18 C night)
in 15.2-cm plastic pots, containing a peat moss-vermiculite
mixture (Jiffy-Mix, Jiffy Products of America, West Chi-
cago, Ill.) mixed with sand (3 :1 v/v). and were watered daily
with Hoagland's nutrient solution.
Ethylene Treatment. The general technique of treating the
third true leaf of intact 3-week-old cotton plants with air, and
the rest of the plant with ethylene, or vice versa, is illustrated
in Figure 1. Small Lucite chambers consisting of two equal
halves were fitted together over the third true leaf blade and
sealed with a gasket of closed cell neoprene 0.64 cm thick. A
slit cut in the gasket allowed the petiole to pass directly through
the center of the gasket. Lanolin was used to make this seal
air tight and elastic bands were placed around the leaf cham-
Plant Physiol. Vol.

E
F-~.0
55, 1975

4
ETHYLENE AND ABSCISSION

FIG. 1. Chamber arrangement for maintaining the third

petiole and abscission zone, to ethylene.

arrangement and the appropriate gas flow equipment, the

true

ber to hold the two halves of the leaf chamber firmly together
against the gasket material. The plants were then placed into
a larger Lucite chamber of 320 liter capacity. Using this
isolated leaf blade could be exposed to either air or ethylene
while the rest of the plant received the same or the opposite
treatment.
Two large Lucite chambers, each accommodating up to 16
plants were used in this study. The internal environmental
conditions in these chambers were 1800 ft-c; 14-hr photo-
period; relative humidity 76% day, and 50% night; tempera-
leaf blade of cotton in air while

O,

-LJ
CfW 6 0
80

40

02
.tiN <

C2 4
C2H4
323

exposing the rest of the plant, including the

ture 28.5 C day, 18 C night. Gas flow through the leaf cham-
bers was maintained at a constant rate of 300 cc/min while
flow through the larger chamber was held constant at 18 1/min.
The inlet and outlet ports on the small chambers were located
at opposite corners. Air or ethylene (14 jul/l) was pumped into
a manifold located inside the large chamber which distributed
the gases to the various flow meters regulating the flow into
each leaf chamber. A slight negative pressure was maintained
on the exit manifold to facilitate the removal of the effluent
air. New tubing was used in each experiment to avoid ethylene
contamination problems. The flow design was such that
ethylene from the same source could be used to treat isolated
leaf blades and entire plants simultaneously, thereby ensuring
that the differences observed were not due to differences in
ethylene concentration. The concentration of ethylene in the
air was monitored throughout each experiment.
Auxin Transport. The classical donor-receiver agar cylinder
technique (10) was used to measure auxin transport capacity
in 5-mm petiole sections. The donor agar cylinders contained
2 ftM naphthaleneacetic acid-i-"4C with a radiochemical purity
of 98.2% as determined by paper chromatography (20) and
liquid scintillation counting (19). Petiole sections were cut
from the distal portion of the petiole 1 cm below the leaf
blade. Sections were allowed to transport auxin for 4 hr in the
dark at 30 C. The sections were then cut in half, placed directly
into a dioxane scintillator fluid (26), extracted overnight on a
00
2 3 4 5 6 7
DAYS
FIG. 2. Effect of ethylene (14 ul/l) on abscission of the third
true leaf of cotton under conditions where the entire plant is ex-
posed to the gas or under conditions where the third true leaf blade
is maintained in air while the rest of the plant is exposed to the gas.
shaker at 4 C, and counted. Donor and receiver agar cylinders
were assayed in a similar manner.
Naphthaleneacetic Acid Leaf Treatments. Leaf blades of the
third true leaf ot 3-week-old cotton plants were immersed
twice each day in 0, 1, 10, 100, 1000 ,uM naphthaleneacetic acid
solutions containing 0.01 % Tween 20. Plants were con-
tinuously exposed to 14 .1/1 of ethylene for 10 days except for
brief periods when the plants were removed from ethylene for
treatments.
RESULTS
Ethylene was totally incapable of inducing abscission of the
third true leaf, even after 7 days, when the leaf blade was
maintained in air (Fig. 2, lower curve). These results were
in marked contrast to the rapid abscission which occurred
when the whole leaf was exposed to ethylene (Fig. 2, upper
curve). Under these conditions 20% of the third true leaves

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324
Table I. Comparisoni of Au.xini Tranisport Capacity in Petiole of
Thiird Leaf of Cottonz tunlder Various Treatmenzt Contditiotis
Inivolviiig Air anid Ethylene
Transport Capacity'
Treatment Condition
2 Days 6 Days
Entire plant in air 52 47
Entire4 plant in C2H4, 14 ,Al/l 11
Leaf blade in air, rest of leaf and plant 30 19
in C2H4
Per cent transport capacity
cpm in basal segment + receiver X 100
cpm in section + receiver
Also see_reference 9 and Fig. 2.
had abscised by the end of the 1st day with 100% abscission
having occurred by the 3rd day. By the end of the 5th day
all of the other leaves on these plants had abscised (data not
shown). As reported for cotton (29), ethylene caused the
younger leaves to abscise first. The older, fully expanded, first
true leaf was the last leaf to abscise. Keeping the third true leaf
blade in air had no significant effect on the abscission rate of
the other leaves on the plant, because by the end of the 5th
day all of these leaves had also abscised. Exposing the third
true leaf blade to ethylene with or without the leaf chamber
had no effect on the rate of abscission (data not shown).
Five such experiments of this nature were conducted for a
maximum length of 12 days. After being purged with air
7 to 12 days in the leaf chambers, the leaf blade began to
senesce. Shortly after visible senescence was observed, the
leaf abscised. Occasionally after 7 days an abscission zone
would form at a point on the petiole just outside the leaf blade
chamber. The formation of an abscission zone in a region
where it does not normally occur is of interest in light of
previous studies where a similar phenomenon has been ob-
served (14, 37, 38).
The auxin transport capacity in the petiole of the third true
leaf under various treatment conditions is presented in Table
I. As reported (9), a 2-day ethylene exposure of the entire leaf
reduced auxin transport capacity in the petiole by 79% when
compared to the control (entire plant in air). Keeping the leaf
blade in air reduced the effectiveness of ethylene in terms of
transport inhibition but it did not completely prevent this effect
of the gas. Auxin transport was reduced 79% when the entire
leaf was treated with ethylene and 42% when the leaf blade
was maintained in air. Under the latter condition auxin trans-
port capacity continued to decline and reached 60% inhibition
by the 6th day. Auxin transport capacity could not be deter-
mined at the end of 6 days in the petioles of the third true
leaves of plants where the entire leaf was in ethylene, since
these leaves had all abscised by the end of the 3rd day.
To determine if the abscission preventive effect of main-
taining the leaf blade in air was related to auxin flow from
the leaf blade to the abscission zone, the auxin transport in-
hibitor DPX18402 (8) was applied as a lanolin ring to the
petiole (Fig. 3). The left-hand portion of Figure 3 again shows
the preventive effect of keeping the leaf blade only in air.
These petioles were treated with plain lanolin and served as
ai DPX1840: 3,3a-dihydro-2-(p-methoxyphenyl)-8H-
pyrazolo [5, 1-a] isoindol-8-one.
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/
BEYER Plant Physiol. Vol. 55, 1975
the controls. The right-hand portion of Figure 3 shows the
effect of applying a lanolin ring containing 100 ,ug/ g of
DPX1840 (about 50 mg) around the petiole 1.5 cm below the
leaf blade at the time the plants were placed into ethylene.
DPX1840 applied in this manner completely nullified the
abscission preventive effect of keeping the leaf blade in air.
After DPX1840 treatment the air-treated leaf blades abscised
just as rapidly as those in ethylene. These rather dramatic
results strongly suggested that auxin was involved in the
preventive effect of keeping the leaf blade in air.
The results presented in Figure 3 also confirm the reported
(8, 28) synergistic effect of DPX1840 with ethylene in abscis-
sion. When the entire plant was treated with ethylene without
DPX1840 application, 33% of the leaves abscised by the
end of the 1st day. Adding DPX1840 to the petiole under the
same conditions increased the abscission percentage obtained
on the 1st day to 68%. DPX1840 applied to the third true leaf
of cotton without added ethylene did not cause abscission
(data not shown).
The data presented above, especially the DPX1840 data,
strongly suggested that the abscission preventive effect of
keeping the leaf blade in air was due to the prevention of an
essential abscission requiring auxin-ethylene interaction in the
leaf blade. Since blocking transport in the petiole with
DPX1840 overcame the effect of keeping the leaf blade in
air, it appeared that the essential ethylene function was to
reduce the flow of auxin out of the leaf blade. If this hypothesis
was correct, then the application of a synthetic auxin to the
leaf blade in the presence of ethylene should also prevent
abscission.
To test this idea naphthaleneacetic acid was applied at
various concentrations as a dip treatment to only the third true
leaf blade (Fig. 4). As previously described (9, and Figs. 2 and
3) ethylene induced rapid abscission of the third true leaf
blade reaching 100% abscission in the experiment by the
end of the 2nd day. Increasing the concentration of naph-
thaleneacetic acid from 1 tuM to 1 mM resulted in a progres-
sively greater reduction of ethylene-induced abscission. At 1
and 10 /IM all of the third true leaves eventually abscised,
but at 0.1 mM less than one-half of these leaves abscised in
10 days and at 1 mm none of them abscised. At the higher
naphthaleneacetic acid concentrations, abscission of the first
and second true leaves was also reduced but not as dramatically
as that of the treated third true leaf.
Since the action of ethylene on the leaf blade is an absolute
requirement if rapid abscission is to occur, it is of interest to
z-100-
c
cm
o 80-
Y.
60-
Q
8 40-
2
C.. 20-
DAYS DAYS
FIG. 3. Effect of DPX1840 on the rate of ethylene (14 ul/1) in-
duced abscission of the third true leaf of cotton when applied as a
lanolin ring to the petiole under conditions where the entire plant
is exposed to the gas or under conditions where the third true leaf
blade is maintained in air while the rest of the plant is exposed to
the gas.
Plant Physiol. Vol. 55, 1975 ETHYLENE AND ABSCISSION 325

the leaf blade continues unaltered ethylene cannot induce

100 / abscission. If this interpretation is correct, then blocking the
/ flow of auxin from the leaf to the abscission zone with an
o< 80 coN'Th auxin transport inhibitor should result in rapid abscission. As
U) J 15M NAA the data presented in Figure 3 demonstrate, the potent in-
CO)-i hibitor of auxin transport DPX1840 (8) does in fact restore
U .a
60' ethylene sensitivity to the leaf. DPX1 840 treatment completely
I NAA
w A negated the preventive effect of keeping the leaf blade in air.
z-
WQ- 40 _
a Additional support for this idea is provided by the data pre-
0
sented in Figure 4 demonstrating that treatment of only the
A4 third true leaf blade with naphthaleneacetic acid under condi-
20 / * I4 MA" tions where the entire plant was exposed to ethylene also
prevented abscission. Naphthaleneacetic acid clearly blocks
I0
1_31
oALE
2 3 4 5 6 7 8 9 10
the effect(s) of ethylene in the leaf blade just as it does in the
petiole and abscission zone (9, 27).
DAYS IN C2H4 (14,XL/L) It is well established (9, 11) that ethylene at the concentra-
FIG. 4. Effect of ethylene (14 IAI/I) on abscission of the third tion used in this study severely reduces auxin transport in the
true leaf of cotton under conditions where the leaf blade is treated petiole of cotton within 24 hr. Therefore, the question arises
with increasing concentrations of naphthaleneacetic acid (NAA) as to why ethylene did not fulfill the same role as DPX1840
and the entire plant is exposed to ethylene. Each datum point is the since the petiole was exposed to the gas for 7 days. The data
average per cent abscission of the treated third true leaf of 10
plants.

ask whether or not this effect alone can trigger abscission. To 100 . ._.
answer this question the leaf blade of the third true leaf was
treated with 14 ,ul/l of ethylene while the rest of the plant was
exposed to air. As indicated by the data presented in Figure 5, Z LL

exposure of only the leaf blade to ethylene does not induce

abscission during a 5-day exposure period. Only when the leaf
blade, petiole, and abscission zone receive ethylene does rapid
abscission occur.
Treating only the leaf blade with ethylene resulted in a
significant decline in the auxin transport capacity even though
the petiole itself was never exposed to the gas (Table II). In
fact, the degree of inhibition after 2 days was essentially the
same as that observed in similar leaves where the whole plant
was treated with ethylene. Transport capacity was reduced
F. S~~~
80% when the whole leaf received ethylene and 82% when
only the leaf blade was treated with ethylene. The degree of
transport inhibition did not change significantly after the
2nd day. 4

DAYS

DISCUSSION FIG. 5. Effect of ethylene (14 Csl/4)on abscission of the third

true leaf of cotton under conditions where the entire plant is ex-
Considerable evidence (1, 3, 16, 32) suggests that a cellular posed to the gas or under conditions where the third true leaf blade
aging process must occur in the cells of the separation region is treated with ethylene while the rest of the plant is maintained in
before they become fully responsive to the abscission stimulat- air.
ing effects of ethylene. Changes in cell sensitivity have largely
been attributed to changes in the auxin concentration of the Table II. Comparison of Auxin Transport Capacity in Petiole of
cells in this region (7, 16). Several workers (9, 11, 13) have Thlird True Leaf of Cotton unlder Various Treatment
suggested that ethylene participates in lowering the auxin con- Conditions Involving Air anld Ethylene
centration in the cell separation region and, hence, in chang-
ing the sensitivity of these cells, through its reported effects Transport Capacity'
on auxin transport (9, 11), destruction (22, 31), and synthesis Treatment Condition
(36). Of these three, the effect of ethylene on auxin transport 2 Days 4 Days
has been most closely correlated with the abscission process.
In the present study the leaf blade was isolated from ethylene
to ascertain whether or not this tissue is a potential site of Entire plant in air 56 52
ethylene action in abscission. If no change in the rate or Entire plant in CX4, 14 IAI/I 11
extent of abscission is observed under such conditions, then Leaf blade in C2H4, rest of leaf and 10 8
ethylene obviously performs no essential function in this plant in air
tissue. However, as the data presented in Figure 2 indicate,
this is not the case. Isolating the leaf blade from ethylene Per cent transport capacity
prevented abscission, clearly demonstrating that treatment of cpm in basal segment + receiver
only the abscission zone and petiole with ethylene is not suffi-
cient to induce rapid abscission. cpm in section + receiver
These results suggest that as long as the flow of auxin from Also see reference 9 and Fig. 2.

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326 BEYER
presented in Table I, suggest a possible answer to this paradox.
Auxin transport capacity was reduced 79% after 48 hr when
the entire leaf was treated with ethylene but only 42% when
the leaf blade remained in air. By keeping the leaf blade in
air the petiole was presumably provided with auxin, which,
as previously reported (9, 33), reduces the ability of ethylene to
inhibit auxin transport. In contrast to ethylene, DPX1840
inhibited auxin transport over 90% within 4 hr under all treat-
ment conditions (data not shown). This marked difference
between the extent and rapidity of transport inhibition by
ethylene and DPX1840 would appear to explain the inability
of ethylene to cause abscission the same as DPX1840 under
these conditions.
Noteworthy with regard to the relative amount of auxin
which might be expected to reach the abscission zone under
conditions where the leaf blade was held in air is a comparison
of this amount with that expected to reach the abscission zone
when the whole leaf is exposed to ethylene. Spectrofluorometric
determinations (35) of IAA levels in the leaf blade following
a 48-hr exposure of the whole leaf to 14 ,ul/l of ethylene
indicate that IAA levels are reduced by 65% (unpublished
data). Following the same period of time auxin transport is
inhibited 79% (Table I). If 100 units of IAA were transported
through the petiole per unit of time, prior to ethylene exposure,
then following a 48-hr ethylene treatment the amount of IAA
would be reduced to only 7 units (65% reduction in the amount
of auxin in the leaf plus a 79% reduction in auxin transport).
On the other hand, when the leaf blade is maintained in air
the amount of IAA would only be reduced to 58 units since no
change in IAA levels of the leaf blade occurs (unpublished
data) and transport is reduced only by 42% under these con-
ditions. While this type of assessment is admittedly over-
simplified, it does serve to illustrate that a 7-fold difference in
auxin content at the abscission zone under these two treatment
conditions could easily occur.
Ethylene apparently alters auxin synthesis, conjugation,
or destruction in the leaf blade, since the total extractable
auxin content, as indicated above, is reduced by 65% following
a 48-hr exposure to ethylene (unpublished data). Effects on
synthesis are suggested by the results of two studies (18, 36)
indicating that ethylene inhibits the conversion of tryptophan
to IAA in Coleus while effects on destruction are suggested
by the enhanced IAA oxidase activities observed in cotton
leaves following ethylene treatment (22). Added to these
potential auxin lowering actions of ethylene is the observed
fact that ethylene reduces auxin transport in the veinal tissues
of the leaf blade in essentially the same manneras it does in the
petiole (unpublished data). Thus, reduced auxin transport and
synthesis, plus enhanced destruction, may contribute to the
over-all decline in auxin efflux from the leaf blade.
Exposure of the petiole and abscission zone to ethylene,
but not the leaf blade. does not result in abscission (Fig. 2).
Similarly. exposure of only the leaf blade to ethylene also
does not cause abscission (Fig. 5), even though auxin transport
is severely inhibited (TableII). This is not too surprising since
the localized application of potent auxin transport inhibitors to
the petiole also does not trigger abscission in the absence of
applied ethylene. Presumably, there is a lack of abscission
under these conditions because there is insufficient ethylene at
the abscission zone to trigger enzyme induction (2, 23, 34) and
secretion (5). This view is supported by the observation that
the sensitivity of leaves to ethylene is increased by pretreating
only the leaf blade with ethylene. When leaf blades of the
third true leaf were pretreated for 48 hr with 14 il/l of
ethylene and then the entire plants were placed in 3 1d/l of
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Plant Physiol. Vol. 55, 1975
ethylene, the leaves abscised, on the average, 1 day earlier than
those which were not pretreated with ethylene.
The results obtained in this study indicate that rapid defolia-
tion of cotton by ethylene involves an essential auxin lowering
function in the leaf blade. This effect coupled with an ethylene-
mediated auxin transport inhibition in the petiole quickly drops
the concentration of the auxin at the abscission zone to a point
where the cells in this region become responsive to the gas in
terms of enzyme induction (2, 23, 34) and secretion (5), lead-
ing to leaf drop. All of these ethylene-mediated processes are
readily reversible since the removal of ethylene at any time up
to the time of actual cell separation halts the abscission process
(9, 17).
In contrast to the role of ethylene in the petiole and abscis-
sion zone, the role of the ethylene in the leaf blade during
natural abscission is not presently clear. Whether or not the
increase in ethylene production during senescence (11, 24)
contributes significantly to the natural decline in auxin con-
tent is not known. The answer to this question must await
further clarification as to the exact nature of those processes
in the leaf involved in this decline plus an assessment of the
sensitivity of these processes to ethylene.
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