Journal of Horticultural Science

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Effects of ethephon on macadamia racemes

R. A. Stephenson & E. C. Gallagher

To cite this article: R. A. Stephenson & E. C. Gallagher (1987) Effects of ethephon on macadamia
racemes, Journal of Horticultural Science, 62:4, 539-544, DOI: 10.1080/14620316.1987.11515818

To link to this article: https://doi.org/10.1080/14620316.1987.11515818

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Journal of Horticultural Science (1987) 62 (4) 539-544

Effects of ethephon on macadamia racemes

By R. A. STEPHENSON and E. C. GALLAGHER
Maroochy Horticultural Research Station, Queensland Department of Primary Industries, P.O.
Box 5083, Sunshine Coast Mail Centre, Nambour, Queensland 4560 Australia

SUMMARY
Ethephon sprays applied to Macadamia integrifolia trees caused the abortion of elongat-
ing racemes. The critical concentration of ethephon for early raceme damage was
between 400 and 800 mg 1· 1• Dormant racemes were less sensitive, and rapidly elongating
racemes more sensitive to ethephon damage. Raceme and floret death and abscission
occurred between two and five weeks after application, and a second flush of racemes
subsequently occurred four to seven weeks after application. The final numbers of
racemes produced were similar, regardless of ethephon concentration. Initial nut set was
reduced but more nuts were set on later racemes. Final nut set and yield were unaffected
by ethephon treatments. This apparent compensating mechanism may have practical
application in crop management strategies.

FLORAL initiation in macadamia occurs before MATERIALS AND METHODS

the nuts from the previous season are mature
(Stephenson, 1980). Since ethylene can acceler-
ate floral senescence and death (Abeles, 1973;
Weaver, 1972) ethephon sprays applied to aid
harvest nut drop may adversely affect sub-
sequent flowering. In fact, Kadman and Ben-
Tal (1983) reported the complete failure of
spring flowering after high concentrations of
ethephon (2000 mg 1·') had been applied to
macadamias at harvest. Adverse effects were
not observed when lower concentrations of eth-
ephon ( <1000 mg J· 1) were applied early,
before raceme elongation. Nakata (1976) also
reported reduced flowering after ethephon
applications.
Gallagher and Stephenson (1986) discussed
the advantages of using ethephon on mac-
adamias in April/May when the nuts are mature
to aid harvesting. Indeed, many large commer-
cial growers in Australia are already doing this.
It was observed however that, when appli-
cations were delayed until June, abscission of
early developing racemes occurred. It is there-
fore important to establish the effects of harvest
season applications, particularly those applied
late, on flowering and subsequent yield. This
paper also examines the time required for eth-
ephon to induce raceme abortion and the sus-
ceptibility of racemes at different stages of
development to ethephon.
Experimental trees were mature, eight to
ten-year-old macadamia trees growing in a
commercial orchard (Experiment 1) and at the
Maroochy Horticultural Research Station
(Experiments 2 and 3), both near Nambour
(Lat. 26° 37's and <50 m above sea level). The
progression of floral development in this
environment is: floral initiation in early May; a
period of raceme dormancy (induced by low
temperature, Moncur et at., 1985) usually
extending to mid-July; raceme elongation
during July and August; anthesis in mid-Sep-
tember and nut set in late September/early
October. In Experiment 1, macadamia trees (cv
Own Choice), on which early June applications
of ethephon had caused the abscission of early
developing racemes, were used to study the
effects on later inflorescence development and
yield. Five representative sub-sample branches
1 em in diameter at the base and initially bear-
ing similar numbers of racemes, were selected
on each of three replicate trees which had been
previously hand sprayed to run off with 0, 400,
800 or 1600 mg J· 1 ethephon plus 0.25 ml J· 1
Agral60 (wetting agent) on a clear, 23°C day as
described by Stephenson and Gallagher (1986).
The pH of the solutions were not adjusted. A
preliminary experiment showed that pH ad just-
men! with 0.25% borax (pH 6.0) had no effect
on the efficacy of ethephon (pH 2.8 at
540 Effects of ethephon on macadamia racemes
1600 mg )·').The racemes were categorized into
developing ( <60 mm long) and developed
{>60 mm) and the numbers of each recorded at
four weekly intervals until the period of peak
anthesis in September a'nd again in December
after premature nut drop had ceased.
Observations to determine the time taken for
ethephon to induce raceme abscission were
carried out on equivalent Own Choice trees not
previously treated with ethephon (Experiment
2). Elongating racemes at 50 tagged nodes on
representative sub-sample branches were hand
sprayed to run-off with 1600 mg J·' ethephon
plus 0.25 mil·' Agral60 wetting agent on a clear,
16°C day in early August. A similar number of
comparable nodes with untreated racemes
were also tagged. Racemes numbers were
recorded at the time of treatment and at two to
four weekly intervals until they stabilized (peak
anthesis) and again after nut set.
Experiment 3 was conducted to determine
the stage(s) of raceme development most sensi-
tive to different concentrations of ethephon.
Designated stages of raceme development
were: dormant ( <5 mm long); developing
(>5 mm, but floral buds not expanded); green
bud (racemes and individual floral buds fully
expanded but unopened); \ind white bud
(anthesis). Treatments were hand-sprayed to
run-off during clear, 16° to 18°C days in July-
August on sub-sample branches containing
racemes at the appropriate stage of develop-
ment on two replicate trees of the cv HAES 333
(Ikaika). Racemes other than those at the stage
of development required were removed prior
to treatment. Ethephon concentrations were 0,
100, 200, 400, 800, 1200, 1600 and 2000 mg J·'
plus 0.25 mJJ·' Agra160. Solution pH was not
adjusted. Racemes were counted and obser-
vations of their condition made at 0, 3 and 5
weeks after treatment and again in late January
(pre-harvest) when nuts borne on remaining
racemes were also counted.
RESULTS AND DISCUSSION
Ethephon applied to induce harvest drop of
nuts in June also induced abscission of early
developed (old) racemes (Experiment 1,
Figure 1). At 400 mg J·' early racemes were
reduced by about 60%, four weeks after treat-
ment (Figure 1B), compared with control
racemes (Figure 1A). At higher con-
centrations, few early racemes survived after
four weeks (Figures 1C and 10). This is consis-
tent with the general effect of ethylene in pro-
moting rapid senescence and death of flowers
(Abeles, 1973).
The observed sequence of ethephon damage
to racemes was: rapid tip necrosis, death and
abscission of florets and abscission of the rachis
which commenced two weeks after treatment
and continued over a four to five week period.
A second flush of racemes occurred within a
month and was most intense where 400 to
800 mg J·' ethephon was applied. On control
trees the appearance of this second flush was
delayed (Figure 1A) and the number of
racemes was lower than on branches sprayed
with 400-800 mg J·' of ethephon (Figures 1B and
1C).
In macadamia there are three buds in the axil
of each leaf. The one closest to the petiole is
most developed, and the third one, least devel-
oped. It is therefore possible for nine inflores-
cences to be produced per node in M.
integrifolia, but this seldom occurs. Usually
only one or two develop at each node. It seems
reasonable that the remaining dormant buds at
a node should be capable of developing
inflorescences to replace these if they are
destroyed at an early stage.
Ethephon, particularly at 400 mg J·', stimu-
lated the development of greater numbers of
racemes during the second flush. In pineapple,
ethylene is used to induce floral initiation
(Abeles, 1973). Trochoulias (pers. comm,
1983) also noted that ethephon applied to mac-
adamia trees in April, immediately before the
stage of floral initiation in early May (Moncur et
al., 1985), promoted greater numbers of floral
buds which appeared earlier. The effect on
macadamia may be through increased floral ini-
tiation or breaking dormancy of preformed
floral tissue (Abeles, 1973). Macadamia trees
may have a compensatory mechanism to main-
tain flowering level, particularly when racemes
are destroyed early during development and
growth. Recent observations showed that a
second flush of macadamia racemes replaced
those destroyed by early frost (S. Newett, pers.
comm., 1985). Such responses may enable use-
ful cultural manipulations to be made. It should
be possible, for example, to compact the har-
 R. A. STEPHENSON and E. C. GALLAGHER 541

B. 400 mg 1-• ethephon

20 A. control •
New
•
Total

10
•
Old

I
LSD IP<0.051
I
.s;;
<.>
"
.0"'
a;
c.
"'Q;
.c
E
:J

" 20
C. 800 mg 1-• ethephon D 1600 mg 1-• ethephon
"
E
"
<.>
"'
a:

10

0 -
I
••
I I I I
•
I
• • • •
4 8 12 16 20 24 4 8 12 16 20 24
Time after spray<ng (weeks I
FIG. 1
Numbers of racemes <+=new, late developing, <60 mm long at 4 weeks after treat-
ment; •=old, early developed, >60 mm long at 4 weeks after treatment; •=total) on
five sub-sample branches of macadamia trees (Own Choice) at 4, 8, 12 and 24 weeks
after they had been sprayed With ethephon in June at (A) 0, (B) 400, (C) 800 or (D)
1600 mg 1· 1 to induce harvest nut drop of the previous crop.

vest season and thus produce savings in pest
control and harvesting. Futhermore, control of
the reproductive development cycle may lead
to improved efficiency in the utilization of
assimilate resources and higher yields.
The initial effects of ethephon of flowering
appeared to be damaging, but the similarity of
total numbers of racemes remaining, regardless
of treatment, at the nut set stage (Figure 1)
indicate that this is not necessarily so. In fact,
total numbers of nut set on sub-sample
branches were similar for all treatments
(7.3±1.5 nuts). In an unpublished raceme
removal study, 0, 25, 50 and 75% of all racemes
were removed from each of four mature, repli-
cate trees (J. Stock, pers. comm., 1985), and
premature nut drop and the number of mature
nuts at harvest were unaffected by the treat-
ments but nut number per raceme was signifi-
cantly greater after 75% of the racemes had
been stripped from trees. Consequently, the
ultimate effect of early adverse occurrences is
often relatively minor either through the pro-
duction of additonal racemes or a greater reten-
tion of nuts per raceme at harvest.
Most of the nuts on control branches (97%)
and those treated with 400 mg P ethephon
(79%) were borne on the early racemes (first
flush) compared with only 13% or Jess on
branches treated at higher concentrations.
Urata (1954) also reported that initial nut set
was highest from early flowering. It therefore
appears that the second flush of flowers serves
to compensate for losses during the earlier
flowering flush. Macadamias characteristically
produce a great excess of racemes and flowers,
few of which set nuts (Urata, 1954; Ito, 1980;
Sakai and Nagao, 1985). Whole-tree yields for
all treatments were similar (22.7±2.9 kg per
tree), thus confirming that the flowering set-
back caused by ethephon was temporary and
that the tree has the capacity to compensate for
earlier losses. There were no apparent residual
effects of ethephon treatments on the yield of
these trees in the following seasons. Yields
were similar to those in the year of treatment.
542 Effects of ethephon on macadamia racemes
When ethephon was applied to developing
(elongating) macadamia racemes from the
second (main) flush, there was no apparent
effect on raceme abortion up to two weeks later
(Experiment 2, Figure 2). Over the next five
weeks, however, death and abscission of
racemes occurred rapidly. The few remaining
racemes (14) set a total of 34 nuts, compared
with 80 nuts on 52 control racemes. In this case,
ethephon was applied so late that there was
little scope for the trees to compensate for
raceme losses and the number of nuts set was
substantially less. Even so, the fewer racemes
set more nuts per raceme than the controls
which is consistent with the experience from
other raceme removal studies.
In Experiment 3 dormant racemes were rela-
tively resistant to ethephon sprays (Figure 3A),
whereas racemes at later stages of development
were sensitive, particularly at concentrations
greater than 400 mg J·' (Figure 3B, C and D).
Raceme survival after exposure of dormant
floral buds (Figure 3A) declined steadily over
the long floral development period (May to
September) and was similar to that on control
branches at the pre-harvest stage. This is con-
sistent with the usual behaviour of macadamia
in setting and retaining nuts from only 0.3% or
less of the flowers produced (Ito, 1980).
Although growing racemes (greater than
100
80
.._
Q)
..c
E 60
::J
c:
Q)
~
u
40
co
0:
20
0
I I I ephon in generating ethylene is dependent on
0 2 4 6 8 10 12 14
Time after ethephon treatment
(weeks)
FIG. 2
Observations of raceme survival on untreated branches of
Own Choice macadamia trees (e) and those hand sprayed
<•).
with 1600 mg 1· 1 ethephon Datum points represent the
total number of racemes at each of 50 tagged nodes.
5 mm long, Figure 3B) were initially insensitive
to ethephon at 400 mg 1- 1 or less, the usual com-
mercial concentrations of 200 to 400 mg 1·'
caused a substantial drop in raceme survival
five weeks later. Thus, although abscission of
the rachis had not occurred after three weeks,
many individual floral buds had aborted and
damage was evident. At 800 mg J·' or more,
substantial ethephon damage to racemes
occurred soon after treatment. By the pre-
harvest stage, however, raceme survival was
similar for all treatments, again reflecting the
usual small proportion of nuts set compared
with flowers produced.
Fully developed racemes (Figure 3C), and
particularly those at anthesis (Figure 3D) tend
to be sensitive to ethephon damage, even at
lower concentrations, and this became obvious
within three weeks of treatment. Thereafter,
little change in relative raceme survival
occurred. At least some racemes sprayed with
low concentrations of ethephon, however,
retained nuts through to the pre-harvest stage.
Overall there was a tendency for nut number
per branch to decrease with increasing con-
centration of ethephon (6.5, 5.0, 4.5 and 4.0
nuts at 0, 100, 200 and 400 mgt-' respectively).
No nuts were produced at concentrations
higher than 400 mg )· 1• These data indicate that
the critical concentration for severe damage
was 400 mg ]·'.
Although concentrations greater than
800 mg ]· 1 damaged racemes severely, the
effects were not as devasting as those reported
by Kadman and Ben-Tal (1983). They found
that, at 2000 mg ]·', but not at lower con-
centrations, ethephon was responsible for the
complete failure of macadamia flowering in
spring. Different environmental conditions in
Israel may underlie these varying responses of
macadamia to ethephon. Several authors
(Biddle et al.; Klein et al., 1979; Olien and
Bukovac, 1978; Swzyjewicz and Kliewer, 1982)
have demonstrated that the efficacy of eth-
temperature and humidity as well as on the pH
of the spray solution. In the field, inconsistent
'responses to ethephon were attributed to differ-
ences in climate and irrigation regimes (Klein et
al., 1978). Abeles (1973) concluded that,
because of such variables, the duration of effec-
tive ethylene concentration is difficult to deter-
 R. A. STEPHENSON and E. C. GALLAGHER 543

Time after treatment· 3 weeks 5 weeks Pre-harvest

A. dormant Inflorescence buds (<5 mr'n long)

B grow1ng racemes (>5 mm long)

15
>
"'c. 1.0
Cll
c:
o_
-g_ -g 0.5
~ §
Q).E
Q; ~
.:::: "'
0 ~'-----
"' !:;
OlQ)
C fully developed racemes (>60 mm long)
E~ 1.5
·m u
E ;;;
~?ft. 10
Cll-
Q)
E
~ 0.5
"'
a:
0

D racemes at anthes1s

II I I I I Ill I I I I I II I I I I I I
01 2 4 8 12 01 2 4 8 12 16 20 01 2 4 8 12 16 20
Ethephon spray concentration (x 100 mg 1" 1 )
FIG. 3
The effect on raceme survival at 3 and 5 weeks after treatment and prior to harvest (m late January) of ethephon
concentratiOns from 0 to 2000 mg I·', applied to: (A) dormant raceme buds ( <5 mm long); (B) developing
racemes (>5 mm long); (C) green bud, fully elongated racemes (>60 mm long) with developed unopened green
tloral buds; and (D) white bud/anthesis (tlower buds openmg) Datum pmnts are the means of 2 subsample
branches (arc-smc transformed). Bars represent the LSD (P <0.05) for each stage of raceme development.

tive ethylene concentration is difficult to deter-
mine. Hence the interpretation of results from
field studies is also difficult.
The commercial practice of applying moder-
ate concentrations ( <400 mg !-')of ethephon to
induce harvest drop of nuts is unlikely to
damage the subsequent crop under Australian
conditions so long as applications are not
delayed beyond the stage when racemes are
just starting to elongate (Experiment 1). Dor-
mant racemes are relatively insensitive to eth-
ephon and, although elongating racemes may
be damaged, the tree has the capacity to
develop a second flush of racemes (Figure 1)
which can compensate for the loss of early
racemes. If ethephon sprays are delayed
beyond this stage, however, the capacity to
compensate for losses is more limited, apart
from an increase in number of nuts retained per
raceme (Experiment 2).
Because of the sensitivity of elongating
racemes to ethephon, it may be feasible to use
544 Effects of ethephon on macadamia racemes

this chemical as a flower-thinning agent to con-
trol crop load, to increase the efficiency of tree
resource utilization and perhaps to optimize
nut quality. Data in Figure 3 indicate that con-
centration of 400 mg )· 1 or less, applied as
racemes commence elongation, may be suit-
able for this purpose by reducing competition
from excess racemes without the danger of
destroying all racemes and thus complete crop
failure.
This work indicates that there is scope for
manipulating the reproductive cycle of mac-
adamia to advantage with ethephon.
We thank Mr T. Grant for access to trees for
this work. Support from Ciba-Geigy in supply
Ethrel (R) and Plus 50 (R) wetting agent is
gratefully acknowledged.

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(Accepte4 17 February 1987)