CSIRO PUBLISHING

Functional Plant Biology, 2012, 39, 481492

http://dx.doi.org/10.1071/FP11254

Time of pruning affects fruit abscission, stem carbohydrates

and yield of macadamia
Lisa McFadyen A,C, David Robertson A, Margaret Sedgley B, Paul Kristiansen B
and Trevor Olesen A,B
A

NSW Department of Primary Industries, 1243 Bruxner Highway, Wollongbar, NSW 2477, Australia.
The University of New England, Faculty of Arts and Sciences, Armidale, NSW 2351, Australia.
C
Corresponding author. Email: lisa.mcfadyen@dpi.nsw.gov.au
B

Abstract. Macadamia (Macadamia integrifolia Maiden and Betche, M. tetraphylla Johnson and hybrids) orchards in
Australia are typically hedged around anthesis (September). Such hedging reduces yields, largely through competition for
carbohydrates between early fruit set and the post-pruning vegetative ush, but also through a reduction in photosynthetic
capacity caused by the loss of canopy. We examined whether hedging at other times might mitigate yield losses. Hedging
time was found to affect yields across four cultivars: A4, A38, 344 and 816. Yield losses were lower for trees hedged
in NovemberDecember than for trees hedged in September. Yields for trees hedged in June were higher than for trees
hedged in September in one experiment, but were similar in a second experiment. Yield losses for September and October
hedging were similar. Hedging time changed the pattern of uctuations in stem water-soluble carbohydrates (WSC). WSC
declined shortly after hedging in September, October or November, and the declines preceded increases in fruit abscission
relative to unpruned control trees. The increase in fruit abscission was less pronounced for the trees hedged in November,
consistent with the idea that fruit become less sensitive to carbon limitation as they mature.
Additional keywords: canopy management, fruit set, hedging, shoot growth.
Received 11 November 2011, accepted 9 April 2012, published online 28 May 2012

Introduction
Macadamia (Macadamia integrifolia Maiden and Betche,
M. tetraphylla Johnson and hybrids) is a recurrent ushing
evergreen tree grown commercially in the subtropics for its
edible kernel. Flowers are borne on racemes, usually initiated
in leaf axils at ~40140 cm from the branch tip depending on
cultivar and season (Salter et al. 2005). In Australian orchards, the
trees are regularly side-hedged for orchard ventilation to facilitate
harvest and to control pathogens, for light penetration for
groundcover growth to mitigate soil erosion, and for
machinery access and effective spraying.
In Australia, hedging typically occurs around anthesis in the
spring (September) after harvest has been completed. However,
hedging at this time accentuates fruit-drop, leading to lower
yields. The accentuated fruit-drop seems to be primarily
caused by competition for carbohydrates between the young
fruit and the post-hedging shoot growth, with a secondary
effect being related to the loss of the photosynthetic canopy,
which presumably reduces the size of the available carbohydrate
pool (McFadyen et al. 2011).
The sensitivity of young fruit to competition and carbohydrate
availability is common amongst horticultural tree species
(Quinlan and Preston 1971; Doud and Ferree 1980; Yuan and
Huang 1988; Berter and Droz 1991; Mehouachi et al. 1995;
Gmez-Cadenas et al. 2000). The fruit appear to become less
Journal compilation
CSIRO 2012

sensitive later in the season as they increase in size. In apple

(Malus domestica Borkh.), shading and girdling treatments
designed to reduce carbohydrate supply to the developing fruit
do not increase fruit abscission when applied after early fruit-drop
(Berter and Droz 1991).
This raises the possibility that hedging macadamia trees
much earlier or later than anthesis, thereby avoiding strong
growth of new shoots during the fruit-drop period, may reduce
the yield penalty associated with hedging.
We compared September hedging with hedging in October,
November, December and June, and with unpruned control
trees. October hedging stimulates the development of a new
ush around late fruit set. November and December hedging
delay the post-pruning vegetative ush until after fruit set.
June hedging allows the post-pruning ush to expand before
fruit set, so that the new ush is a source of carbon to the
young fruit. June hedging may also promote owering by
removing the immature ush that may inhibit bud release
(Olesen 2005) and by synchronising bud release with
conditions that favour oral development (Olesen et al. 2002).
Hedging time may also affect the production of fruiting wood
in subsequent seasons as more short shoots are produced
under higher temperatures (Wilkie et al. 2009a) and short
shoots are more likely to ower than long shoots (Wilkie et al.
2009b).
www.publish.csiro.au/journals/fpb

482

Functional Plant Biology

We conducted three experiments using four commercial

cultivars: A4, A38, 344 and 816 (Peace et al. 2002).
The cultivars A4 and 344 represent the extremes of the
oral habit of macadamia, with A4 bearing its racemes close
to the tips of the major branches and 344 bearing its racemes
much deeper within the canopy (Salter et al. 2005). A38 and
816 are similar to 344 but do not bear their racemes quite
so deep within the canopy. A range of oral habits was chosen
because earlier work (McFadyen et al. 2011) showed that
competition between ush development and fruit set was
greater the closer the racemes were to the new shoots; it may
be that cultivars that bear their racemes nearer to the edge of the
canopy are more susceptible to competition effects.
To investigate the effect of hedging time on production, we
measured: whole-tree owering, fruit-drop or set, and yield. To
explore the potential link between fruit drop/set and carbon
availability we monitored changes over time in both fruit
numbers on selected racemes and the concentrations of stem
water-soluble carbohydrates (WSC). In addition, we measured
the numbers and lengths of post-pruning shoots on selected
branches to assess the potential for hedging time to affect
fruiting wood production in subsequent seasons.
Materials and methods
All experiments were conducted in commercial orchards in
northern New South Wales, near Alstonville (28.9S,
153.5E), Australia on deep, well drained reddish-brown clay
loam soils (Morand 1994). Trees were grafted on H2 seedling
rootstock (Macadamia integrifolia Maiden and Betche) and
orchard operations were carried out according to industry
guidelines (OHare et al. 2004).
Experiment 1: the effects of September, October
and December pruning on fruit per raceme, yield
and shoot growth in cultivars 344 and A4
This experiment was conducted on 344 (M. integrifolia) and
A4 (M. integrifolia  M. tetraphylla Johnson) trees at two
sites: Brooklet (28.7S, 153.5E) and Alstonville.At Brooklet,
the trees were 10 years old and those at Alstonville were 17 years
old. At each site, the cultivars were in separate but nearby
plantings and were planted in northsouth rows. Treatments
included two pruning times, September to early October
(anthesis) and December (1013 weeks after anthesis (AA)),
and an unpruned control. At Brooklet, trees were tip-pruned using
a motorised hedge trimmer on a long shaft, removing at least the
most recent ush from most branches. At Alstonville, trees were
pruned with a commercial hedging machine. Two cuts were made
on each side of the hedgerow: a vertical cut at the bottom of the
canopy and a second cut above this, angled slightly towards the
centre of the row. Hedging removed ~30 cm of canopy from each
side of the hedgerow.
The experiments were laid out as randomised complete blocks
in ve and three rows for 344 and A4 trees, respectively, at
Brooklet, and in two and three rows for 344 and A4 trees,
respectively, at Alstonville. For each experiment, there were six
blocks, each comprising three plots of three trees, and treatments
were allocated randomly to plots within blocks.

L. McFadyen et al.

The 344 trees were pruned on 26 September 2007 and 11

December 2007 (10 weeks AA) at Brooklet, and on 4 September
2007 (anthesis) and 6 December 2007 (13 weeks AA) at
Alstonville. The rst pruning time at Brooklet had been
planned to coincide with anthesis but owering occurred over
an extended period and it was not possible to time pruning to
coincide with a clear peak. At the time of pruning, there were
racemes with green unopened orets, racemes at anthesis and
racemes at advanced sepal fall in approximately equal numbers.
The A4 trees were pruned on 5 October (anthesis) and 12
December (10 weeks AA) at Brooklet, and 26 September
(anthesis) and 6 December (10 weeks AA) at Alstonville.
Fruit retention, shoot number and length of the post-pruning
ush were measured at the Alstonville site. Fruit per raceme
were counted on 10 tagged racemes per tree on one tree per plot
on 11 October 2007 (5 weeks AA) and 5 November 2007 (6 weeks
AA) for 344 and A4, respectively. Counts were made on
three trees per plot on 8 November 2007 (9 weeks AA) and 11
February 2008 (23 weeks AA) for 344, and on 3 December 2007
(10 weeks AA) and 13 February 2008 (20 weeks AA) for A4.
After pruning, 10 pruned branches were tagged on each tree, and
shoot number and length were measured when the shoots were
fully elongated.
At both sites, fruit were harvested from the ground between
March and July 2008 at 4-week intervals. Fruit were dehusked
and samples were taken from each plot at each harvest to
determine moisture content. Yield was expressed as nut-inshell at 10% moisture content, which is the commercial
standard.
Experiment 2: the effects of June and September pruning
on owering, fruit abscission, yield and shoot growth
in cultivars 816 and A38
The planting used for the experiment was located at Knockrow
(28.7S, 153.5E) and comprised alternate rows of 8-year-old
816 and A38 (M. integrifolia) trees planted in northsouth
rows spaced at 6 m with 5 m and 3 m within-row spacings for
816 and A38 trees, respectively. Average tree heights were 6.3
and 5.7 m for 816 and A38, respectively. Trees were sprayed
with 2-chlorethylphosphonic acid (Ethrel, Bayer CropScience)
at 600 parts per million on 11 May 2005 to promote fruit
abscission (Trueman et al. 2002) so that harvesting could be
completed before the June pruning.
Trees were pruned on 78 June 2005 (14 weeks before
anthesis) or at anthesis on 1214 September 2005. A
motorised hedge trimmer mounted on a long shaft was used to
tip-prune the sides of the trees up to a height of 4.5 m. Tippruning involved the removal of at least the most recent ush
from all branches up to this height. Each pruning time  cultivar
combination was replicated four times in a randomised
complete block design over four rows. Blocks comprised two
plots of each cultivar, which each comprised three and four
trees for 816 and A38, respectively.
After pruning, 10 branches were tagged on each tree. Shoot
number and length on each branch were measured between 3
and 4 weeks after the start of shoot growth. Shoot length
measurements were repeated ~1-week intervals until the
shoots had nished elongating. The daily mean temperature,

Pruning time, growth and yield in macadamia

calculated as the average of the daily maximum and minimum

temperatures, was sourced from records at the Centre for Tropical
Horticulture, Alstonville, and used in the analysis of shoot
growth.
Raceme density on each tree was visually assessed on 27
September 2005 using a scale from 1 to 5, with 1 and 5
representing the lowest and highest raceme densities,
respectively.
Fruitlets shed from the tree were collected from the orchard
oor from each plot and weighed on 8 November 2005. A
subsample of 100 fruits from each plot was also weighed and
used to estimate individual fruit weight; this was then used with
the total fruit weight per plot to estimate fruit number per plot.
Mature fruit were harvested from the orchard oor on four
occasions for each cultivar between March and September 2006
at 4 to 5-week intervals. Fruit were dehusked and samples of
nuts were taken at each harvest to determine moisture content.
Yield was expressed as nut-in-shell at 10% moisture content.
On the second and third harvests, which together yielded more
than 95% of the total harvest, samples of 100 nuts were taken
from each plot to estimate nut moisture and individual nut
weight. The latter was used with the total nut weight per plot
to estimate nut number per plot. Samples were not collected
from individual plots at the other harvests as there were too few
fruit. Instead, a composite sample was collected from the four
plots for each pruning time  cultivar combination to estimate
moisture content.
Experiment 3: the effects of June, September, October
and November pruning on owering, fruit per raceme,
yield, shoot growth and stem carbohydrates in cultivar A4
This experiment was carried out on 7-year-old A4 trees at
Lindendale (28.9S, 153.4E). Fifty trees over ve rows were
divided into 10 blocks of ve trees. Treatments comprised four
pruning times and an unpruned control arranged in a complete
block design. The allocation of trees to treatments was based on
the yields from the previous season (hand-harvested in May
2006), with the trees restrictively randomised within blocks
such that across the blocks, the treatments had similar
pretreatment yield distributions. Trees were tip-pruned in 2006
with secateurs to remove at least the most recently matured
ush from most branches. Pruning dates were 6 June
(~4 months before anthesis), 18 September (anthesis), 16
October (4 weeks AA) or 8 November (7 weeks AA).
Racemes per tree were counted near anthesis, and racemes
per tree with fruit were counted just before harvest. Ten racemes
on each tree were tagged near anthesis and the numbers of fruit
per raceme were counted weekly from 3 to 13 weeks AA, then
fortnightly to 19 weeks AA and then a month later at 23 weeks
AA.
Fruit were stripped from the trees on 3 April 2007. Fruit
were dehusked and samples of 100 nuts were taken from the
harvest from each tree to determine moisture content and
individual nut weight. Total nut weight for each tree was
expressed at 10% moisture content.
Stem samples for carbohydrate analyses were collected at
intervals from ve replicates of each treatment. Samples were
rst collected from June-pruned and control trees on 16 June

Functional Plant Biology

483

2006, ~2 weeks after pruning, and again on 18 July and 3 August

2006, then weekly until 17 January 2007. Collection from the
other treatments commenced on 7 September 2006, ~2 weeks
before anthesis, and again on 28 September 2006 and then weekly
until 17 January 2007. Sample collection and preparation for
carbohydrate analysis followed the procedure described by
McFadyen et al. (2011), which produced a ground sample of
bark and wood for analysis. The samples were analysed for WSC,
which make up more than 80% of total-non-structural
carbohydrates in macadamia (Olesen et al. 2008). WSC were
extracted by incubating 100 mg of powder in 20 mL 0.2% w/w
benzoic acid solution with gentle agitation at room temperature
for 1 h; then spinning out the residue at 945g. The supernatant
was analysed using the anthrone method (Yemm and Willis
1954). Five millilitres of the anthrone reagent (760 mL sulfuric
acid, 330 mL of water, 1 g of thiourea and 1 g of anthrone) were
added to a frozen 1-mL sample of the supernatant while the
sample was shaken in a vortex mixer. This mixture was then
incubated in a boiling water bath for 3 min, transferred to an icewater bath for 10 min and returned to room temperature before
being measured in a spectrophotometer at 620 nm in a 10-mm
cuvette. The standards were solutions of 0500 mg L1 D(+)glucose in 0.2% w/w benzoic acid.
New shoots around the tip-pruning cut were collected from
one branch per tree on the dates stems were collected for
carbohydrate analyses to estimate the timing and rate of postpruning shoot growth. Shoots were dried to a constant weight at
60C. The daily mean temperature, calculated as the average of
the daily maximum and minimum temperatures, was sourced
from records at the Centre for Tropical Horticulture, Alstonville,
and used in the analysis of shoot growth.
Statistical analyses
Linear mixed models (ASReml-R software; Butler et al. 2007)
were used to analyse pruning treatment, cultivar, site and time
effects on raceme, fruit, nut, stem WSC and some shoot data.
A cubic spline term for time was included as a random effect in
analyses of fruit number per raceme and stem WSC. F-ratio tests
(Wald method) and likelihood ratio tests were conducted to test
the statistical signicance of the xed and random effects,
respectively. The LSD was used to compare means.
Non-linear regressions (Sigmastat, Systat Software, Inc., San
Jose, CA, USA) were used to describe changes in shoot length
and dry weight (DW) over time. The function tted to treatment
means in Figs 1 and 2 was y = a (1 + eb(c x))1, where y is shoot
length or DW and x is time, and a, b and c are parameters. The
maximum increase in shoot length or DW = 0.25  a  b and
occurs at x = c. The function was also tted to individual replicates
and ANOVAs were carried out on the estimates of maximum
rate of shoot length or DW increase, and the average of the
mean daily temperature for the 3 days around c.
Results
Experiment 1: the effects of September, October
and December pruning on fruit per raceme,
yield and shoot growth in cultivars 344 and A4
There was a signicant effect of treatment on yield of 344 trees
(P < 0.05) and no interaction between treatment and site

484

Functional Plant Biology

L. McFadyen et al.

Date of measurement
Aug.
160

Sep.

Oct.

Date of measurement

Nov.

Aug. Sep. Oct. Nov. Dec. Jan. Feb.

Dec.

September

June

15

(a)

June

140
120
100

816

80

A38

10

60
40
20

Early fruit drop

15

10

(b)

September

(c)

October

(d )

November

Weeks before/after anthesis

(P > 0.05). Yield was 18% and 54% higher in December-pruned

trees than in September-pruned trees at the Brooklet
and Alstonville sites, respectively (Table 1). At neither site
was there any difference between December-pruned and
unpruned control trees (Table 1; P > 0.05). For A4 trees,
there was a signicant effect of treatment on yield (P < 0.05),
and a signicant interaction between treatment and site (P < 0.05).
At Brooklet, December-pruned trees had 95% more yield than
September-pruned trees and 15% less than control trees
(Table 1). At Alstonville, there was no difference between
treatments (Table 1; P > 0.05).
Initial fruit set in A4 trees at Alstonville , measured at
56 weeks AA, was very low, at less than two fruit per raceme
and with only 63% of racemes with fruit, compared with around
seven fruit per raceme and 98% of racemes with fruit in 344
(Table 2). There was a tendency for fruit retention to be lower
in September-pruned 344 trees compared with the other 344
treatments (Table 2).
Shoots in the post-pruning ush tended to be shorter in
December-pruned trees than in September-pruned trees,
although not signicantly for 344 (Table 1; P > 0.05). There
was not a consistent effect of pruning time on shoot number. In
344 trees, shoot numbers were lower in December-pruned trees
(P < 0.05), whereas in A4 trees, shoot numbers were lower in
September-pruned trees (P < 0.05).
Experiment 2: the effects of June and September pruning
on fruit abscission, yield, owering and shoot growth
in cultivars 816 and A38
There was a signicant effect of pruning time on yield (P < 0.01)
and no interaction with cultivar (P > 0.05); June-pruned 816 and
A38 trees produced 41% and 16% more yield than September
pruned trees, respectively (Table 3), although the difference
was not signicant for A38 (P > 0.05). There was no effect
of pruning time on raceme density (Table 3). Early shoot growth

10

Shoot dry weight (g)

Fig. 1. The average shoot length of new shoots on branches of 816 and
A38 macadamia trees tip-pruned in June or September, increased
sigmoidally over time. Points are means of four replicates, based on
plots of three and four trees for 816 and A38, respectively, with 10
branches per tree and several shoots per branch (see Table 3). The curves
represent predicted values from the tted non-linear function (P < 0.0001),
y = a (1 + eb(c x))1, where y and x are variables, and a, b and c are
parameters.

15

10

15

10

0
10

10

15

20

Weeks before/after anthesis

Fig. 2. The dry weight of new shoots on branches of A4 macadamia trees
tip-pruned in (a) June, (b) September, (c) October or (d) November in 2006
increased sigmoidally over time. Points are means of ve replicates (trees).
The line represents predicted values from the tted non-linear function
(P < 0.0001), y = a (1 + eb(c x))1, where y and x are variables, and a, b
and c are parameters.

in September-pruned trees coincided with premature fruit

abscission, whereas shoot growth from June-pruned trees had
almost fully elongated before early fruit-drop (Fig. 1). Septemberpruned trees tended to drop more fruit than trees pruned in June
and to have fewer fruit per tree at harvest (Table 3), albeit not
always signicantly.
Final shoot length was 32% and 62% shorter in Septemberpruned 816 and A38trees, respectively (Table 3; P < 0.0001)
but there was no effect of pruning time on shoot number per
branch (Table 3; P > 0.05).

Pruning time, growth and yield in macadamia

Functional Plant Biology

Table 1. Nut weight per tree and shoot length and number for 344 and
A4 macadamia trees pruned in September or December in 2007, or for
unpruned control trees
Values are means and s.e. for six plots of three trees. Within each cultivar,
treatment means followed by a different letter are signicantly different
(P < 0.05)
Site and
treatment

Nut weight
per tree (kg)
344 A4

Shoot length
(mm)
344 A4

Shoot
number
344 A4

Alstonville (17-year-old trees)

Control
11.7b 15.5c
September
7.6c 14.7c
December
11.7b 14.5c

73.2a 123.7a
66.0a 90.0b

5.1a
3.5b

Brooklet (10-year-old trees)

Control
19.5a 16.7a
September
16.1a 7.3b
December
19.0a 14.3c

s.e

3.6

5.7

1.9

0.7

3.1a
3.6b

0.17 0.13

Maximum shoot growth rate was higher in September-pruned

trees than June-pruned trees, corresponding with higher
temperatures in September (P < 0.001 for shoot growth rate
and temperature, data not shown).
Experiment 3: the effects of June, September, October
and November pruning on owering, fruit per raceme,
yield, shoot growth and stem carbohydrates in cultivar A4
Fruit number per raceme declined curvilinearly with time (Fig. 3).
There was a signicant effect of treatment on fruit number per
raceme (P < 0.0001), and a signicant interaction between
treatment and the spline term for time (P < 0.0001). Fruit
numbers per raceme for the spring pruning times and the
control were similar at the start of the sampling period
(P > 0.05) but fruit abscised faster in the pruned treatments,
with fruit number per raceme declining relative to the control
at 5, 6 and 10 weeks AA for the September, October and
November pruning times, respectively (P < 0.05). The start of

Table 2. Fruit per raceme and percentage of racemes with fruit at three different times after anthesis (AA) for 344and A4 macadamia
trees pruned in September or December in 2007, or for unpruned control trees
The results correspond with the Alstonville site yields in Table 1. Values are means and s.e. for six trees at 56 weeks AA and six plots of three
trees for 910 and 2023 weeks AA. Within each cultivar, treatment means followed by a different letter are signicantly different (P < 0.05)
Cultivar and
treatment

56 weeks AA
Fruit per
Percentage
raceme
of racemes
with fruit

344
Control
September
December
s.e.
A4
Control
September
December
s.e

910 weeks AA
Fruit per
Percentage
raceme
of racemes
with fruit

2023 weeks AA
Fruit per
Percentage
raceme
of racemes
with fruit

6.8a
7.0a
6.0a

98a
91a
97a

1.4a
1.1b
1.5a

61a
45b
63a

1.0a
0.8a
1.1a

49a
34a
53a

0.8

0.16

0.17

1.8a
1.3a
1.4a

63a
53a
47a

0.7a
0.4b
0.5ab

36a
27a
29a

0.4a
0.2b
0.2b

24a
18a
17a

0.2

0.12

0.08

Table 3. Raceme density, fruit-drop, fruit number per tree, individual nut weight, nut weight per tree, and shoot length and number
for 816 and A38 macadamia trees pruned in June or September 2005
Values are means and s.e. for four plots of three or four trees for 816 and A38, respectively. Treatment means followed by a different letter
are signicantly different (P < 0.05). Raceme density was visually assessed using a scale from 1 to 5, with 1 and 5 representing the lowest and
highest raceme densities, respectively
Cultivar and
Raceme
pruning time density
816
June
September
A38
June
September
s.e.

485

Number of fruit dropped

Fruit number Individual nut Nut weight
Shoot
Shoot number
in fruit diameter classes
per tree
weight (g)
per tree (kg) length (mm)
per branch
>5.0 mm 2.55.0 mm <2.5 mm

1.9a
2.5a

268a
324ab

163a
291a

6a
14a

1850a
1238b

7.5a
7.8a

15.5a
11.0b

164a
111b

3.5a
3.4a

3.1b
3.2b

460b
660c

933b
987b

76b
72b

1345b
1153b

7.7a
7.7a

11.8b
10.2b

150a
93b

4.0a
3.1a

0.3

52

119

10

115

0.15

0.9

0.4

486

Functional Plant Biology

L. McFadyen et al.

Date of measurement
Oct.

Nov.

Dec.

Jan.

Feb.

Mar.

Oct.

Nov.

Dec.

Jan.

Feb.

Mar.

100.0
50.0

(a)

Control

(b)

June

(c)

September

(d)

October

25.0
12.0
6.0
3.0

Fruit per raceme +1 (log scale)

1.5
1.0
100.0
50.0
25.0
12.0
6.0
3.0
1.5
1.0
100.0

November

(e)

50.0

Control
June
September
October
November

(f)

25.0
12.0
6.0
3.0
1.5
1.0
0

10

15

20

10

15

20

25

Weeks after anthesis

Fig. 3. Fruit number per raceme over time in A4 macadamia trees that were (a) unpruned, or on which branches were
tip-pruned in (b) June, (c) September, (d) October or (e) November 2006. Dashed lines in (ae) are the observed means of
10 racemes per tree. Solid lines are predicted values from a linear mixed model. The predicted lines for (ae) are shown
together in (f). There was a signicant effect of treatment (P < 0.0001) and a signicant interaction with a spline term for
time (P < 0.0001).

the relative decline in fruit number in October- and Novemberpruned trees occurred before early shoot growth, which,
in those treatments, occurred at around 7 and 9 weeks AA,
respectively (Fig. 2). Shoot growth in the Septemberpruned trees commenced at around 5 weeks AA (Fig. 2),
which coincided with the decline in fruit number relative to
the control.
The fruit abscission rate in the control trees slowed slightly
in early November and then increased a few weeks later, with
fruit continuing to drop at this rate until 18 weeks AA in late
January, after which there was little further abscission (Fig. 3).
The fruit-drop period nished earlier in the spring-pruned
treatments compared with the control, with little abscission
occurring after 11, 13 and 16 weeks AA in the September-,
October- and November-pruned trees.

The percentage of racemes with fruit at maturity and fruit

number per tree were both lower in the September- and Octoberpruned trees relative to November-pruned trees and the
control (Table 4; P < 0.05). There were no differences in the
percentage of racemes with fruit or fruit number per tree between
the November-pruned trees and the control, or between the
September- and October-pruned trees (Table 4; P > 0.05).
The effects on fruit retention were reected in yield, with
November-pruned trees yielding 77% and 152% more than
Septemberand
October-pruned
trees,
respectively
(Table 4; P < 0.05). There were no signicant differences
between September- and October-pruned trees or between
November-pruned trees and the control (Table 4; P > 0.05).
June pruning resulted in a substantial reduction in raceme
numbers per tree (Table 4; P < 0.05) to around 50% of that in

Pruning time, growth and yield in macadamia

Functional Plant Biology

487

Table 4. Raceme number at anthesis, percent of racemes with fruit at maturity, fruit number per tree, individual nut weight and nut weight per tree for
A4 macadamia trees that were unpruned (control) or on which all branches were tip-pruned in June, September, October or November 2006
Values are means and s.e. for 10 trees. Treatment means followed by a different letter are signicantly different (P < 0.05)
Treatment

Raceme number per tree

at anthesisA

Control
June
September
October
November

6.98a
6.30b
7.06a
6.88a
6.93a

s.e

0.16

Fruit number
per tree

Individual nut
weight (g)

Nut weight
per tree (kg)

38a
18b
16b
15b
32a

756a
206b
270b
228b
622a

8.8a
9.6b
10.1b
8.7a
7.7c

6.5a
2.1b
2.7b
1.9b
4.8a

74

0.16

0.6

Values for raceme number per tree at anthesis are means and s.e. for log-transformed data, with back-transformed values in brackets.

the other treatments. There were also far fewer fruit per
raceme at the start of the sampling period than in the other
treatments (Fig. 3; P < 0.05). The percentage of racemes with
fruit, fruit number and yield for June-pruned trees were all
substantially lower than in November-pruned trees
and control trees (Table 4; P < 0.05), and were not
signicantly different from September- or October-pruned
trees (Table 4; P > 0.05).
June pruning also reduced stem WSC during winter (Fig. 4).
There was a signicant effect of treatment (P < 0.01), and a
signicant interaction between treatment and time (P < 0.01).
Stem WSC rose and fell in both the control and June-pruned
trees, but rose less and fell more in the June-pruned trees. Stem
WSC in the June pruned trees was lower than in the control trees
from 10 weeks before anthesis to near anthesis in September,
when stem WSC was around half that in the control trees (Fig. 4;
P < 0.05).
Stem WSC also rose and fell during the springsummer
period in all treatments (Fig. 5). There was a signicant effect
of treatment (P < 0.05) and a signicant interaction between
treatment and the spline term for time (P < 0.0001). Stem
WSC for all treatments rose from early September and then
declined following a peak in mid-October. WSC in the spring
pruning times and the control were similar at the start of
sampling (P > 0.05), but rose less in September-pruned trees
compared with the control and fell more in the October- and
November-pruned trees. Consequently, the September-,
October- and November-pruned trees had lower WSC than
the control from 27, 510 and 913 weeks AA, respectively
(P < 0.05). The start of the decline in stem WSC in each
spring pruning treatment relative to the control preceded the
increase in fruit-drop in that treatment relative to the control.
It also preceded early post-pruning shoot growth in each
treatment (Fig. 2).
Although June-pruned trees had lower WSC in early
September, levels increased to a higher mid-October peak than
in the other treatments (Fig. 5; P < 0.05).
There was a third clear peak in WSC in the September- and
October-pruned trees in December, and a much smaller peak
around this time in the control but not in the November- or Junepruned trees (Fig. 5).
The rate of shoot DW increase did not correspond with
temperature. Shoot growth rate was highest in September-pruned

Date of measurement
June
10

July

Aug.

Sep.

Control

(a)

8
6
4
2
10

WSC (% d.wt)

(1080)
(46)
(1169)
(970)
(1024)

Percentage of racemes
with fruit at maturity

June

(b)

8
6
4
2
10

Control
June

(c)

8
6
4
2

14

12

10

Weeks before anthesis

Fig. 4. Dawn levels of stem water-soluble carbohydrates (WSC) over time,
in A4 macadamia trees on which branches were (a) unpruned or (b) tippruned in June. Dashed lines in (a) and (b) represent observed values for one
sample per tree. Solid lines are predicted values from a linear mixed model.
The predicted lines for (a) and (b) are shown together in (c) There was a
signicant effect of treatment (P < 0.01) and a signicant interaction with a
spline term for time (P < 0.01).

trees, although temperatures were higher in October and

November (P < 0.05, for shoot growth rate and temperature, data
not shown).

488

Functional Plant Biology

L. McFadyen et al.

Date of measurement
Sep.

Oct.

Nov.

Dec.

Jan.

Feb.

Sep.

Oct.

Nov.

Dec.

Jan.

Feb.

10

Control

(a)

(b)

June

(d )

October

September

WSC (% d.wt)

(c)
8

X Axis 2

November

(e)

Control
June
September
October
November

(f )

2
0

10

15

20

10

15

20

Weeks after anthesis

Fig. 5. Dawn levels of stem water-soluble carbohydrates (WSC) over time, in A4 macadamia trees on which
branches were (a) unpruned, or tip-pruned in (b) June, (c) September, (d) October or (e) November 2006. Dashed lines in
(ae) represent observed values for one sample per tree. Solid lines are predicted values, using a linear mixed model. The
predicted lines for (ae) are shown together in (f). There was a signicant effect of treatment (P < 0.05) and a signicant
interaction with a spline term for time (P < 0.0001).

Discussion
Effects of hedging time on fruit abscission, yield,
owering and stem carbohydrates
There was a tendency for less fruit abscission and higher yields
for trees pruned at times that avoided shoot growth during the
early fruit abscission period (Fig. 3; Tables 14). For trees pruned
in NovemberDecember, yields for 7- and 10-year-old A4 trees
were, respectively, 77% and 95% higher than for Septemberpruned trees (Experiments 1 and 3, Tables 1, 4). In 10- and 17year-old 344 trees, yields were 18% and 54% higher,
respectively, in December-pruned trees compared with
September-pruned trees (Experiment 1, Table 1). This is
consistent with the hypothesis that fruit growth is less sensitive
to a reduction in carbohydrate availability when it occurs towards

the end of, or after, the premature fruit drop period. This view was
also supported by the patterns in fruit-drop and carbohydrate
levels observed in trees pruned in September, October and
November in Experiment 3. Pruning reduced carbohydrate
levels and fruit set in all pruning treatments relative to the
unpruned control; however, in the November-pruned trees, the
reduction occurred at the end of the normal fruit-drop period and
the reduction in fruit set was less than in the other pruning
treatments (Figs 3, 5; Table 4). In apple, Berter and Droz
(1991) noted a similar resistance to abscission in fruit when a
disruption to assimilate supply was imposed after the early
summer drop period. They attributed this to greater fruit sugar
levels at this growth stage, which is consistent with observations
in Citrus that low sugar concentrations in young fruit were
associated with increased levels of plant growth regulators that

Pruning time, growth and yield in macadamia

promote abscission (Gmez-Cadenas et al. 2000). Resistance to

abscission is also linked with an increase in auxin supply to the
abscission zone from actively growing distal tissues (Taylor and
Whitelaw 2001). In macadamia, the reduction in fruit abscission
generally observed at around 10 weeks AA coincides with an
increase in fruit growth rate (Trueman and Turnbull 1994;
McConchie et al. 1996).
An exception to the trend for higher yield for November- and
December-pruned trees occurred in the 17-year-old A4 trees in
Experiment 1 (Table 1). This may have been due to low initial fruit
set, which, at 56 weeks, was around half that of 344 trees at
the same site (Table 2) and similar to nal fruit set in A4 trees in
Experiment 3 (Fig. 3, Table 4) and the results found in an earlier
study (McFadyen et al. 2011). Low initial set may have
minimised inter-fruit competition, with the result that the
remaining fruit were less sensitive to competition from postpruning shoot growth.
Trees pruned in June in Experiment 2 also tended to drop fewer
fruit and produce higher yields than trees pruned in September
(Table 3). This was probably because the post-pruning shoot
growth of June-pruned trees had almost fully elongated by
anthesis and was supporting the developing crop, not
competing with it. In lychee (Litchi chinensis Sonn.), pruning
at a time that allowed one vegetative ush to develop before
owering resulted in higher yields than pruning at a later time that
did not allow regeneration of the canopy before owering
(Menzel et al. 2000).
In Experiment 2, there was no effect of pruning time on
owering (Table 3); however, in Experiment 3, June pruning
substantially reduced raceme production to around half that of
the control and the other pruning treatments (Table 4), possibly
due to a large decrease in stem carbohydrates over winter in Junepruned trees relative to control trees. (Fig. 4). Intensity of
owering has frequently been correlated with levels of tree
carbohydrate reserves, although not consistently (Chaikiattiyos
et al. 1994; Goldschmidt 1999; Wilkie et al. 2008). In macadamia,
trunk girdling, which raises carbohydrate availability above the
girdle, increased raceme production (Nagao and Sakai 1990).
In contrast, Wilkie (2009) found differences in return owering
for a range of crop loads despite an absence of differences in stem
carbohydrates close to the time of oral initiation. Although
carbohydrate is unlikely to be the only factor affecting
owering, it is likely that a minimum level is required for bud
release and oral differentiation (Goldschmidt 1999). Another
possible explanation for the low raceme production in Junepruned trees is that pruning increased light intensity to a point
where it inhibited oral initiation. Raceme production in
macadamia is lowest in high light conditions (Olesen et al.
2011) and high light intensity also reduced oral initiation in
olive (Olea europaea L.) (Stutte and Martin 1986) and in the
ornamental tree Metrosideros excelsa Soland. ex Gaertn.
(Henriod et al. 2003). The difference between the two
experiments may relate to differences in canopy density. No
estimates were made, but the tree canopies in Experiment 3 were
visibly less dense than canopies in the other experiments in this
study and in previous studies (McFadyen et al. 2011). This may
have resulted in a bigger impact of leaf area removal on
carbohydrate availability and canopy irradiance. The effect of
June pruning on owering in Experiment 3 was also in contrast to

Functional Plant Biology

489

a series of three experiments in which pruning in late May or

early June did not reduce raceme production (Wilkie et al. 2010).
Despite a lower raceme load for June-pruned trees in
Experiment 3 than for the other treatments, initial fruit set was
also lower (Fig. 3). Generally, fruit set tends to increase as ower
density decreases (Wilkie 2009). It may be that carbohydrates
were reduced to the point where they also limited initial fruit set.
There was no difference in overall fruit abscission or yield
between trees pruned in October at 4 weeks AA and trees
pruned in September (Experiment 3; Fig. 3, Table 4). We had
anticipated that the October timing would minimise any effect
on fruit abscission by avoiding a reduction in stem carbohydrate
during the premature fruit abscission period. This was based on
several reports that fruit-drop in unpruned trees ceased by around
10 weeks AA (Sakai and Nagao 1985; Trueman and Turnbull
1994; McFadyen et al. 2011) and on our earlier study, in which
post-pruning shoot growth and leaf area removal did not affect
fruit abscission until 6 and 8 weeks after pruning, respectively
(McFadyen et al. 2011). However, in the study reported here,
stem carbohydrate and fruit abscission declined relative to the
control within 2 weeks of the October and November pruning
times (Figs 3, 5). This was before shoot growth had commenced
(Fig. 2) and indicates that leaf area removal by pruning had an
earlier effect on assimilate supply than previously reported. This
may have been due to the low canopy density of the trees in this
study, as already mentioned. A lower initial leaf area would have
increased the impact of leaf area removal on assimilate
availability within the tree compared with trees with denser
canopies. Additionally, the fruit-drop period in our study
extended to 18 weeks AA, well beyond the 10 weeks AA
previously reported. This may have allowed more time for the
October pruning to increase fruit abscission. Again, the extended
drop period may have been due to the lower canopy density of the
trees. It was unlikely to be a seasonal effect, as nearby trees of the
same cultivar, in the same season, exhibited a shorter drop period
(McFadyen et al. 2011).
Varietal effects
The response to time of hedging was demonstrated for four
cultivars including A4, a hybrid (M. integrifolia 
M. tetraphylla) that bears owers around 40 cm from the end
of the branch and hence close to the site of the post-pruning ush,
as well as M. integrifolia cultivars 816, A38 and 344, which
bear owers between 80140 cm from the branch apex (Salter
et al. 2005). In our earlier work, the effect of post-pruning shoot
growth on fruit abscission and yield was quite local, with shoot
growth on branches in one part of the canopy having little effect on
fruit abscission on unpruned branches in another part of the
canopy (McFadyen et al. 2011). Although resources do not
appear to move between major branches in response to shoot
growth, our study suggests that they move signicant distances
along a branch to support shoot growth at the expense of fruit
growth, given that all cultivars were affected by hedging time. The
response tended to be greater in the A4 experiments than in
the 344experiments, which may have been because of the
greater distance between shoot and fruit growth sites in the
latter. It may also have been simply due to site differences,
such as the low canopy density in A4 trees in Experiment 3

490

Functional Plant Biology

and the asynchronous owering in 344 at the Brooklet site

in Experiment 1. There is insufcient information to speculate
whether the proximity of fruit to the end of the branch is a general
feature of hybrid cultivars as, within the 15 cultivars assessed by
Salter et al. (2005), only two were known hybrids.
In 344 trees, there was no difference in fruit retention or yield
between trees pruned in December and unpruned control trees. In
A4 there were trends for November- and December-pruned
trees to have lower yields than the control, but not always
signicantly. Given that A4 bears fruit closer to the end of
the branch than 344, it is more likely to have fruit removed
through hedging, and this may result in yield reduction. Our
results suggest that for cultivars that carry fruit well within the
canopy, the negative effect of pruning on yield might be avoided
altogether.
Seasonal variation in WSC and further interactions
with pruning treatments
In general, uctuations in reserve carbohydrates corresponded
with changes in vegetative and reproductive growth, and pruning
at different times substantially varied the location and magnitude
of the peaks and troughs. Stem WSC in control trees increased
during winter (Fig. 4), corresponding with a period when growth
is restricted by low temperature (Moncur et al. 1985; Stephenson
and Cull 1986) and demand for assimilates is low. A similar
accumulation of carbohydrate over winter has previously been
observed for macadamia (Stephenson et al. 1989) and other crops
including avocado (Persea americana Mill.; Scholeeld et al.
1985) and Citrus (Jones and Steinacker 1951). Stem WSC in
control trees peaked in early August then declined until just
before owering in early September (Fig. 4), coinciding with
the start of the late winter to early spring ush (Stephenson and
Cull 1986) and raceme elongation (Moncur et al.1985). There
was a second peak in mid-October (Fig. 5), which was possibly
due to a combination of increased assimilate supply from the
maturing late winter to early spring ush, and the relatively low
rate of fruit growth at that time (Trueman and Turnbull 1994).
The peak may also have been due to an increase in
photosynthesis in response to rising temperature (Lloyd 1991;
Olesen 2011). A similar peak was observed by Wilkie (2009) but
not by Stephenson et al. (1989). The latter study measured
changes in trunk wood carbohydrate, which may respond
differently to changes in carbon supply and demand compared
with stems. Also, the sampling interval of Stephenson et al.
(1989) (monthly) was longer than that used in this study
(weekly) and by Wilkie (2009) (fortnightly), and was therefore
less likely to detect short-term uctuations. From mid-October,
stem carbohydrates in the control trees declined as the rate of
fruit DW accumulation increased (Trueman and Turnbull 1994).
A third, much smaller, peak occurred in mid-December following
an increase in fruit abscission that may have resulted in a
temporary reduction in demand from fruit growth (Figs 3, 5).
A similar recovery around this time, coinciding with increased
fruit abscission, was also observed by McFadyen et al. (2011).
Stem carbohydrate increased again in early January, consistent
with the studies of Stephenson et al. (1989) and Wilkie (2009).
The stem WSC pattern for June-pruned trees over winter was
similar to control trees except that levels increased less in early

L. McFadyen et al.

winter and declined more in the period leading up to anthesis

(Fig. 4). June pruned trees also had a second peak in October but
it was higher than in control trees (Fig. 5). This possibly reected
a greater availability of assimilates due to the synchronised
maturation of the post-pruning ush. Unlike in the control
trees, there was no evidence of a third peak in carbohydrate in
December (Fig. 5); nor was there an increase in fruit abscission
around this time (Fig. 3). This is consistent with our earlier
suggestion that the peak in control trees was related to a
temporary reduction in assimilate demand for fruit growth
following an increase in fruit abscission.
Stem WSC in September-pruned trees rose slightly during
October despite the absence of a recently matured ush (Fig. 5).
This supports the idea that an increase in photosynthesis in
response to rising temperature made some contribution to stem
carbohydrates at this time. October- and November-pruned
trees had similar peaks to the control during October but
declined at a greater rate due to the reduction in photosynthetic
area and the demand for carbohydrates from the post-pruning
ushes (Fig. 5). The September- and October-pruned trees had
larger peaks during December compared with the control trees.
This corresponds to a greater reduction in demand for fruit
growth following accentuated fruit abscission in the pruned
trees (Fig. 3). In addition, increased assimilate supply from the
synchronised maturation of the post-pruning ushes may have
contributed to the rise in stem WSC. The subsequent decline
may have been due to the start of, or an increase in, root growth.
Root growth alternates with shoot growth in a pattern that is
consistent with competition between roots and shoots for
photosynthates, and with inhibition of root growth by auxins
produced in the shoots (Bevington and Castle 1985). The start of
the decline in stem WSC in September- and October-pruned
trees (Fig. 5) coincided with a period of slowing growth in the
post-pruning ushes (Fig. 2).
Effects of hedging time on post-pruning shoot production
Wilkie et al. (2009a) found that higher temperatures decreased
the lengths of post-pruning shoots in non-bearing 849 trees but
increased the number of shoots per branch. In our experiments,
the temperatures in June were cooler than in September, which
in turn were cooler than in October and November (data not
shown; see also Wilkie et al. (2010) and Olesen (2011) for more
general temperature trends). We also found a tendency for
decreased shoot lengths at higher temperatures in 816,
A38, 344 and A4 trees, but no consistent trend in shoot
numbers, with numbers increasing, decreasing or remaining
relatively constant (Tables 1, 3).
Our experiments were on bearing trees, and it may be that
some of the differences in responses between the studies were
related to competition between shoots and developing fruit in
our experiments. Effects of crop load on shoot development
have been previously demonstrated in several crops, including
macadamia (McFadyen et al. 2005; Wilkie 2009), apple
(Palmer 1992), Citrus (Verreynne and Lovatt 2009) and grape
(Vitis vinifera L.; Edson et al. 1993). This might explain why,
for A4 trees, the maximum rate of DW accumulation of new
shoots per branch following pruning in September (early fruit
set; Fig. 2) was greater than the rates following pruning in

Pruning time, growth and yield in macadamia

October or November (fruit ll), contrary to the common

understanding that shoot growth rate reects current
temperature (Keller and Tarara 2010).
The effects of different pruning times on shoot development
may have implications for owering and production in
subsequent years, but the effects need further clarication and
the reproductive behaviours of the various shoots need to be
monitored over many years.
Conclusions
Increased fruit abscission and reduced yield following pruning
was associated with depletion of carbohydrate levels. Pruning
time varied the uctuation pattern of carbohydrate reserves, and
the effect of pruning on fruit abscission and yield was less when
the disruption to carbohydrate supply occurred late in the
premature fruit-drop period; this is consistent with the
hypothesis that fruit are less sensitive to low carbohydrate
levels at later stages of development. Although current
photosynthate is the main source of carbon for growth in
evergreen trees (Olesen et al. 2008), our results suggest that
reserve carbohydrates have an important buffering role,
especially during key phenological events. The difference
between pruning times was demonstrated for cultivars that
bear fruit both near and far from the branch tip, suggesting
that the pruning effect on carbohydrate supply and fruit
retention extended over a signicant distance. The results have
implications for canopy management of macadamia in that the
negative effect of hedging on yield may be minimised by hedging
in June or December rather than the current industry standard of
hedging in September. Shoot length varied with pruning time,
consistent with the idea that shoot length increases with
increasing temperature. However, there was not a consistent
relationship between pruning time and shoot growth rate or
numbers. It is likely that the effect of temperature on these
variables was moderated by crop load.
Acknowledgements
We thank Russell Priddle, Glenn Smith, Steven Muldoon, Alister Janetzki,
Magda Verbeek, Robyn Andrews and Lloyd Morgan for technical assistance,
and Stephen Morris for assistance with and advice on statistical analyses and
R programming and for comments on a draft of the manuscript. We also
thank Cliff James, Greg James, Terry Walker, Surrey Bogg, Lindsay Bryen,
John Townsend and David Rodgers for allowing us access to their orchards,
and for their assistance and cooperation. The work was funded by Horticulture
Australia Limited (project number MC04024) using the Australian
macadamia industry levy and matched funds from the Australian government.

References
Berter J, Droz P (1991) Studies on locating the signal for fruit abscission in
the apple tree. Scientia Horticulturae 46, 201214. doi:10.1016/03044238(91)90043-X
Bevington KB, Castle WS (1985) Annual root growth pattern of young citrus
trees in relation to shoot growth, soil temperature, and soil water content.
Journal of the American Society for Horticultural Science 110, 840845.
Butler D, Cullis BR, Gilmour AR, Gogel BJ (2007) ASReml-R reference
manual. (Queensland Department of Primary Industries and Fisheries:
Brisbane)

Functional Plant Biology

491

Chaikiattiyos S, Menzel CM, Rasmussen TS (1994) Floral induction in

tropical fruit trees: effects of temperature and water supply. Journal of
Horticultural Science 69, 397415.
Doud DS, Ferree DC (1980) Inuence of altered light levels on growth and
fruiting of mature Delicious apple trees. Journal of the American Society
for Horticultural Science 105, 325328.
Edson CE, Howell GS, Flore JA (1993) Inuence of crop load on
photosynthesis and dry matter partitioning of Seyval grapevines.
I. Single leaf and whole vine response pre- and post-harvest. American
Journal of Enology and Viticulture 44, 139147.
Goldschmidt EE (1999) Carbohydrate supply as a critical factor for citrus fruit
development and productivity. HortScience 34, 10201024.
Gmez-Cadenas A, Mehouachi J, Tadeo FR, Primo-Millo E, Talon M (2000)
Hormonal regulation of fruitlet abscission induced by carbohydrate
shortage in citrus. Planta 210, 636643. doi:10.1007/s004250050054
Henriod RE, Jameson PE, Clemens J (2003) Effect of irradiance during oral
induction on oral initiation and subsequent development in buds of
different size in Metrosideros excelsa (Myrtaceae). The Journal of
Horticultural Science & Biotechnology 78, 204212.
Jones WW, Steinacker ML (1951) Seasonal changes in concentration of
sugars and starch in leaves and twigs of citrus trees. Proceedings of the
American Society for Horticultural Science 58, 14.
Keller M, Tarara JM (2010) Warm spring temperatures induce persistent
season-long changes in shoot development in grapevines. Annals of
Botany 106, 131141. doi:10.1093/aob/mcq091
Lloyd J (1991) Modelling stomatal responses to environment in Macadamia
integrifolia. Australian Journal of Plant Physiology 18, 649660.
doi:10.1071/PP9910649
McConchie CA, Meyers NM, Anderson K, Vivian-Smith A, OBrien S,
Richards S (1996) Development and maturation of macadamia nuts in
Australia. In Proceedings of the third Australian Society of Horticultural
Science and the rst Australian Macadamia Society Research
Conference. (Eds RA Stephenson, CW Winks) pp. 234238.
McFadyen LM, Morris SG, McConchie CA, Oldham MA (2005) Effect of
hedging and tree removal on productivity of crowding macadamia
orchards. Australian Journal of Experimental Agriculture 45, 725730.
doi:10.1071/EA04120
McFadyen LM, Robertson D, Sedgley M, Kristiansen P, Olesen T (2011)
Post-pruning shoot growth increases fruit abscission and reduces stem
carbohydrates and yield in macadamia. Annals of Botany 107, 9931001.
doi:10.1093/aob/mcr026
Mehouachi J, Serna D, Zaragoza S, Agusti M, Talon M, Primo-Millo E (1995)
Defoliation increases fruit abscission and reduces carbohydrate levels in
developing fruits and woody tissues of Citrus unshiu. Plant Science 107,
189197. doi:10.1016/0168-9452(95)04111-7
Menzel C, Olesen T, McConchie C, Wiltshire N, Diczbalis Y, Wicks C (2000)
Lychee, longan and rambutan optimising canopy management. Final
report DAQ-177A. (Rural Industries Research and Development
Corporation: Canberra)
Moncur MW, Stephenson RA, Trochoulias T (1985) Floral development of
Macadamia integrifolia Maiden & Betche under Australian conditions.
Scientia Horticulturae 27, 8796. doi:10.1016/0304-4238(85)90058-5
Morand T (1994) Soil landscapes of the Lismore-Ballina 1:100 000 sheet.
Report. (Soil Conservation Service of NSW: Sydney)
Nagao MA, Sakai WS (1990) Effects of gibberellic acid, ethephon or girdling
on the production of racemes in Macadamia integrifolia. Scientia
Horticulturae 43, 4754. doi:10.1016/0304-4238(90)90035-D
OHare P, Quinlan K, Stephenson R, Vock N (2004) Growing guide:
macadamia growers handbook. (Queensland Department of Primary
Industries: Brisbane)
Olesen T (2005) The timing of ush development affects the owering of
avocado (Persea americana) and macadamia (Macadamia integrifolia x
tetraphylla). Australian Journal of Agricultural Research 56, 723729.
doi:10.1071/AR04287

492

Functional Plant Biology

L. McFadyen et al.

Olesen T (2011) Late 20th century warming in a coastal horticultural region

and its effects on tree phenology. New Zealand Journal of Crop and
Horticultural Science 39, 119129. doi:10.1080/01140671.2010.550627
Olesen T, Menzel CM, Wiltshire N, McConchie CA (2002) Flowering and
shoot elongation of lychee in eastern Australia. Australian Journal of
Agricultural Research 53, 977983. doi:10.1071/AR01179
Olesen T, Robertson D, Muldoon S, Meyer R (2008) The role of carbohydrate
reserves in evergreen tree development, with particular reference to
macadamia. Scientia Horticulturae 117, 7377. doi:10.1016/j.
scienta.2008.03.026
Olesen T, Huett D, Smith G (2011) The production of owers, fruit and leafy
shoots in pruned macadamia trees. Functional Plant Biology 38, 327336.
doi:10.1071/FP11011
Palmer JW (1992) Effects of varying crop load on photosynthesis, dry matter
production and partitioning of Crispin/M.27 apple trees. Tree Physiology
11, 1933.
Peace C, Vithanage V, Turnbull C, Carroll BJ (2002) Characterising
macadamia germplasm with codominant radiolabelled DNA
amplication ngerprinting (RAF): markers. Acta Horticulturae 575,
371380.
Quinlan JD, Preston AP (1971) The inuence of shoot competition on fruit
retention and cropping of apple trees. Journal of Horticultural Science 46,
525534.
Sakai WS, Nagao MA (1985) Fruit growth and abscission in Macadamia
integrifolia. Physiologia Plantarum 64, 455460. doi:10.1111/j.13993054.1985.tb08522.x
Salter B, Hardner C, Forrester R, Levitt C, Mathews K, McConchie CA (2005)
Differences in ower position and shoot growth between macadamia
varieties. In Assessment protocols for selection and management in
macadamia. Final report MC00035. (Ed. C Hardner) pp. 126141.
(Horticulture Australia Limited: Sydney)
Scholeeld PB, Sedgley M, Alexander DMcE (1985) Carbohydrate cycling in
relation to shoot growth, oral initiation and development and yield in the
avocado. Scientia Horticulturae 25, 99110. doi:10.1016/0304-4238(85)
90081-0
Stephenson RA, Cull BW (1986) Vegetative ushing patterns of macadamia
trees in south east Queensland. Scientia Horticulturae 30, 5362.
doi:10.1016/0304-4238(86)90081-6
Stephenson RA, Gallagher EC, Rasmussen TS (1989) Effects of growth
manipulation on carbohydrate reserves of macadamia trees. Scientia
Horticulturae 40, 227235. doi:10.1016/0304-4238(89)90115-5

Stutte GW, Martin GC (1986) Effect of light intensity and carbohydrate

reserves on owering in olive. Journal of the American Society for
Horticultural Science 111, 2731.
Taylor JE, Whitelaw CA (2001) Signals in abscission. New Phytologist 151,
323339. doi:10.1046/j.0028-646x.2001.00194.x
Trueman SJ, Turnbull CGN (1994) Fruit set, abscission and dry matter
accumulation on girdled branches of macadamia. Annals of Botany 74,
667674. doi:10.1006/anbo.1994.1169
Trueman SJ, McConchie CA, Turnbull CGN (2002) Ethephon promotion of
crop abscission for unshaken and mechanically shaken macadamia.
Australian Journal of Experimental Agriculture 42, 10011008.
doi:10.1071/EA01164
Verreynne JS, Lovatt CJ (2009) The effect of crop load on budbreak inuences
return bloom in alternate bearing Pixie mandarin. Journal of the
American Society for Horticultural Science 134, 299307.
Wilkie JD (2009) Interactions between the vegetative growth,
owering and yield of macadamia (Macadamia integrifolia,
M. integrifolia  M. tetraphylla) in a canopy management context.
PhD thesis. University of New England, Armidale, Australia.
Wilkie JD, Sedgley M, Olesen T (2008) Regulation of oral initiation in
horticultural trees. Journal of Experimental Botany 59, 32153228.
doi:10.1093/jxb/ern188
Wilkie JD, Sedgley M, Olesen T (2009a) A model of vegetative ush
development and its potential use managing macadamia (Macadamia
integrifolia) tree canopies. Crop and Pasture Science 60, 420426.
doi:10.1071/CP08337
Wilkie JD, Sedgley M, Morris S, Muldoon S, Olesen T (2009b)
Characteristics of owering stems and raceme position in macadamia.
The Journal of Horticultural Science & Biotechnology 84, 387392.
Wilkie JD, Sedgley M, Olesen T (2010) The timing of pruning affects ushing,
owering and yield of macadamia. Crop and Pasture Science 61,
588600. doi:10.1071/CP09167
Yemm EW, Willis AJ (1954) The estimation of carbohydrates in plant extracts
by anthrone. Biochemical Journal 57, 508514.
Yuan R, Huang H (1988) Litchi fruit abscission: its patterns, effect of shading
and relation to endogenous abscisic acid. Scientia Horticulturae 36,
281292. doi:10.1016/0304-4238(88)90063-5

www.publish.csiro.au/journals/fpb