Efforts to understand and improve crop load of ‘Hass’

avocado
H. Hofman1, J.D. Wilkie2, J. Griffin1 and R. Langenbaker1
1Bundaberg Research Facility, Department of Agriculture and Fisheries, Bundaberg, Australia; 2Wollongbar
Primary Industries Institute, Wollongbar, Australia.

Abstract
Avocado (Persea americana Mill.) yields are low compared to other fruit crops,
commonly less than 10 t ha-1 in Australia, prompting our research into improving
productivity through understanding and improving crop load capacity. When we
removed varying percentages of inflorescences on 3-year-old ‘Hass’ trees in a trial at
Childers, central Queensland, Australia, we found clear relationships between number
of inflorescences and yield, yield and vegetative growth, as well as yield and fruit size.
In a second trial, we removed the whole or part of the vegetative tip of indeterminate
inflorescences on 3-(then 4-)year-old ‘Hass’ trees to reduce vegetative competition
with fruit growth. Spring fruit set increased in these tipping treatments in one of two
years of the trial, but final fruit set did not differ from the control in either year. In the
same trial, foliar application of “Cytolin” (®Sumitomo Chemicals, 19 g L-1 6-
benzyladenine and 19 g L-1 gibberellins4+7) at the end of the extension of the spring
flush to increase increasing axillary branching and thus fruiting sites, increased the
total length of summer vegetative growth of shoots but not branching.

Keywords: productivity, vigour, inflorescence, indeterminate, flower thinning

INTRODUCTION
Avocado (Persea americana Mill.) yields are low compared with other fruit crops,
commonly less than 10 t ha-1 in Australia. Wolstenholme (1986) calculated a theoretical
capacity of 32.5 t ha-1 based on energy costs. The relationships between flowering intensity,
fruit set and yield and the factors affecting them are not fully understood, including the
extent to which avocado crop load is affected by competition for resources. There are few
strategies available to producers to increase crop load.
Avocados produce abundant inflorescences in the form of thyrsi, each consisting of
hundreds or thousands of flowers (Chanderbali et al., 2013). However, low levels of fruit set,
followed by heavy fruit drop in spring and summer, result in low yields. The percentages of
fruit set per inflorescence and fruit abscised have been found to be similar from year to year
in several studies, suggesting final yield is strongly dependent on or related to the initial
number of flowers (Dixon et al., 2007; Garner and Lovatt, 2008; Whiley et al., 2013). To test
this relationship, we established a trial on young ‘Hass’ trees in which we thinned varying
percentages of inflorescences to see effects on crop load.
The majority of avocado inflorescences are indeterminate, with a vegetative shoot
extending through the primary axis of the inflorescence, distal to the secondary axes
supporting each cyme of flowers. A minority of inflorescences may be functionally
determinate, with no shoot extension. The vegetative shoot of indeterminate inflorescences
emerges during early fruit development. The competition between vegetative and fruit sinks
during this period appears to be a factor in determining final yield. Management strategies
that adjust this balance, including the use of plant growth regulators that suppress
vegetative growth, such as gibberellin bio-synthesis inhibitors paclobutrazol and
uniconazole, have been shown to increase fruit set (Kohne and Kremer-Kohne, 1987;
Wolstenholme et al., 1990; Gardiazabal et al., 1995; Salazar-Garcia and Lovatt, 2000). Trials
of tip pruning of the vegetative flush in indeterminate inflorescences also increased spring
fruit set (Biran, 1979; Cutting and Bower, 1990). However, some report that gains made in

Acta Hortic. 1228. ISHS 2018. DOI 10.17660/ActaHortic.2018.1228.49 331

Proc. XI Int. Symp. on Integrating Canopy, Rootstock and
Environmental Physiology in Orchard Systems
Ed.: L. Corelli Grappadelli
spring fruit set with these strategies are negated by heavier fruit drop in summer (Cutting
and Bower, 1990; Wolstenholme et al., 1990). This suggests that such strategies need to
ensure sufficient vegetative growth to support ongoing fruit development. We determined to
test a suggestion by Wolstenholme and Whiley (1990) that tipping the vegetative shoot after
three or four new leaves have formed may give a better response than total removal of the
vegetative shoot.
An alternative strategy for increasing crop load may be to increase the number of
fruiting sites. As avocado flowering is predominantly on terminal shoots from the last-grown
flush (Thorp and Sedgley, 1993), this would mean increasing the extent of branching in
summer flushes. Thorp and Sedgley (1993) tested several plant growth regulators on
avocado and found that Cytolin (®Sumitomo Chemicals, 19 g L-1 6-benzyladenine and 19g L-1
gibberellins4+7) applied at 25 mL L-1 increased sylleptic axillary shoot growth and
subsequent fruit set. The timing of application was critical to success: the most successful
treatment date was the end of the extension of the spring flush, when apical buds are
quiescent. Thorp and Sedgley’s work was done with individual shoots: we determined to test
this strategy on whole trees, including a treatment that combined Cytolin application with
vegetative tipping, to maximise potential gains.

MATERIALS AND METHODS

Flowering and crop load trial

At full bloom in 2013 (9-16 September), we applied eight flower thinning treatments
to 2.5-year old ‘Hass’ trees on ‘A8’ rootstock in a commercial orchard at Isis, Queensland,
with single tree plots in four blocked replicates. Treatments were removal of 0 (control), 10,
20, 40, 60, 80, 90 and 95% of inflorescences, both terminal and axillary. The vegetative bud
or shoot of indeterminate inflorescences was not removed. The numbers of inflorescences
tree-1 were counted and converted to inflorescence density (inflorescences m-3 of canopy)
using canopy dimensions measured at full bloom in 2013.
Fruit were harvested at maturity on 1/5/2014 and the individual weight of each fruit
and total fruit weight per tree were recorded. Yield efficiency (kg fruit m-3 of canopy) was
calculated using canopy dimensions measured at the time of harvest.

Tipping and fruiting sites trial

On a block of three-year old ‘Hass’ trees on ‘Velvick’ rootstock in a commercial orchard
near Childers, Queensland, we established a trial of five treatments plus a control (T1)
randomized in six replicates in a single row. Treatments were applied to the same trees over
two consecutive years (2014/15 and 2015/16). In four of the treatments, we removed the
vegetative tip of indeterminate inflorescences when the vegetative shoot was a minimum of
5 cm, either by ‘whole shoot tipping’, that is, removing the whole vegetative portion
including all emerging leaves (Treatment 2) or ‘partial tipping’, that is, removing the tip but
leaving three to five leaves at the base (T3, T4 and T6). We applied treatments either weekly
after vegetative shoots began to elongate (T2 and T3) during the ~six-week period of
flowering and vegetative shoot emergence, or once only at full bloom (T4 and T6). We tipped
every indeterminate shoot on the whole canopy as far as we could reach using 2.8 m ladders
(approximately 80% of the tree). Each shoot was tipped only once. In T5 and T6, we applied
Cytolin as a foliar spray to the whole tree in a single application at the end of the extension of
the spring flush, at a concentration of 15 mL L-1 in the first year and 20 mL L-1 in the second
year. T6 combined the Cytolin application with ‘once only’ partial tipping.
We tagged individual flowering shoots (30 on each tree in Year 1 and 25 in Year 2) to
monitor fruit retention and shoot growth. At the end of the spring and summer flushes, we
counted the number of new branches (axes) and the number of fruit, and measured the total
length of each flush including the length of new branches, on each tagged shoot. ‘Fruiting
sites’ were defined as the number of axes on each shoot. In treatments where the shoot was
not tipped (T1 and T5), this included new branches and the primary axis, but in tipping
treatments (T2, T3, T4 and T6), this included new branches but excluded the primary axis

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because a tipped terminal will not produce an inflorescence in the following spring.
Yields of each tree were weighed at harvest and mean fruit weight per tree calculated
from a random sample of 50 fruit. Yield efficiency was calculated using canopy dimensions
measured at flowering each year.

Data calculation and analysis

Canopy volume (V) was calculated using the formula for an ellipsoid: V = (π×x×y×z)/6;
where x is along row canopy width, y is across row canopy width and z is tree height less the
height of any pruned ‘skirt’. Data were analyzed in Genstat (16th edition), using one-way
analysis of variance for the tipping and fruiting sites trial data and linear and non-linear
regression for the flowering and crop load trial. Means were separated using Fisher’s
protected least significant difference test at α=0.05.

RESULTS

Flowering and crop load trial

Yield tree-1 increased with inflorescence number (Figure 1; r2=0.66, P<0.001) and
yield efficiency (kg m-3) increased with inflorescence density (inflorescences m-3 canopy;
Figure 2; r2=0.59, P<0.001). Mean fruit weight decreased with increasing yield efficiency
(Figure 3; r2=0.53, P<0.001). Number of fruit per inflorescence also decreased with
inflorescence density (Figure 4; r2=0.11, P<0.037). There was also an association between
yield efficiency and vegetative growth, with canopy volume and trunk cross-sectional area at
harvest decreasing with increasing yield efficiency (r2=0.24, P=0.003 (Figure 4) and r2=0.37,
P<0.001 respectively).

Figure 1. The effect of the number of inflorescence tree-1 on yield tree-1 in the flowering and
crop load trial 2013/14.

Figure 2. The effect of inflorescence density on yield efficiency in the flowering and crop
load trial 2013/14.

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Figure 3. The effect of yield efficiency on mean fruit weight in the flowering and crop load
trial 2013/14.

Figure 4. The effect of inflorescence density on fruit per inflorescence remaining at harvest
in the flowering and crop load trial 2013/14.

Topping and fruiting sites trial

The first year of the trial showed no significant differences between treatments in fruit
set on tagged shoots at either the end of spring or summer, or in yield tree-1.
In Year 2, fruit set at the end of spring was significantly higher than the control (T1)
for the weekly partial tipping treatment (T3) and the treatment in which we applied Cytolin
plus a one-off partial tipping (T6) (Table 1). However, by the end of summer there was no
difference in fruit retention between treatments.
In both years, the Cytolin application (T5 and T6) failed to produce a significantly
different mean number of potential fruiting sites compared to the control (T1). The Cytolin
application without tipping (T5) increased mean total shoot length (including the length of
new branches) in Year 2 only, while Cytolin plus tipping (T6) increased total shoot length in
both years. While tipping often resulted in the growth of new branches, the loss of the
fruiting site on the terminal meant that the mean number of potential fruiting sites per shoot
was not significantly different between treatments, except in Year 1 when T2 resulted in
fewer potential fruiting sites (mean of 1.96) than all other treatments including the control
(mean of 3.51) (P=0.032).
There was no significant difference in yield tree-1, yield efficiency or mean fruit weight
between treatments in Year 1 or Year 2 of the trial. There was no correlation on a per tree
basis regardless of treatment between mean number of fruiting sites per shoot that
developed in Year 1 and yield efficiency in Year 2 (r=-0.003, P=0.99).

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Table 1. Fruit set and vegetative growth of tagged shoots and yield efficiency by treatment
in Year 2 (2015/16) of the tipping and fruiting sites trial.
Mean total
Mean
Mean Mean shoot length Mean
end-of-
spring summer at end potential Yield
summer
fruit set fruit drop summer flowering efficiency
fruit set
(fruit (fruit including terminals (kg m-3)
(fruit
shoot-1) shoot-1) new branches shoot-1
shoot-1)
(cm)
T1. Control 0.48ab 0.19ab 0.30 22.7ab 1.72 1.61
T2. Tip whole 0.69bc 0.29abc 0.40 19.0a 1.43 1.40
shoot – weekly
T3. Partial 0.72c 0.40bc 0.33 23.8ab 1.51 1.36
Tipping – weekly
T4. Partial 0.58abc 0.36abc 0.22 23.1ab 1.29 1.50
tipping – once
T5. Cytolin app’n 0.40a 0.16a 0.24 29.6bc 1.65 1.61
T6. Cytolin app’n 0.74c 0.51c 0.23 32.6c 1.76 1.20
+ partial tipping – once
P 0.010 0.029 0.320 0.008 0.250 0.359
s.e. 0.07 0.08 0.06 2.47 0.15 0.15
s.e. = standard error of the means.
1Potential flowering terminals. Includes the originally tagged axis and new branches for treatments 1 and 5 excludes the
originally tagged axis for other treatments as a tipped axis will not produce an inflorescence.

DISCUSSION

Inflorescence density, fruit set and yield

There was a clear increase in yield efficiency with increased inflorescence density in
the flowering and crop load trial (Figure 2). This indicates that flowering intensity can limit
avocado yield, as has previously been identified in relation to ‘off’ years in alternate bearing
cycles (Salazar-Garcia et al., 1998; Dixon et al., 2007; Garner and Lovatt, 2008). At the lower
inflorescence densities, the canopy was able to compensate to some extent by setting more
fruit per inflorescence (Figure 4) and by increasing fruit size (Figure 3), but this was not
sufficient to prevent a reduction in yield efficiency. Similarly, reduced floral bud density in
apple results in greater fruit set per inflorescence and greater fruit size (Breen et al., 2015).
As inflorescence density increased above 30-40 inflorescences m-3 canopy, it is unclear
whether the relationship with yield efficiency continued to increase linearly or whether it
began to plateau (Figure 2). A linear relationship would suggest that yield was sink limited,
with sufficient carbohydrate resources to support increasing numbers of fruit. There is some
evidence within the literature to support this suggestion, with the percentage of avocado
flowers setting and retaining fruit remaining constant regardless of flowering intensity and
whether it is an ‘on’ or ‘off’ year (Dixon et al., 2007; Garner and Lovatt, 2008). A study by
Finazzo et al. (1994) of photoassimilate partitioning during the development of individual
fruiting shoots also suggests that photoassimilates were not limited.
On the other hand, a ‘plateau’ in the relationship between inflorescence density and
yield efficiency would suggest that, as inflorescence density increased above a certain point,
limitations in the supply of carbon assimilates prevented any further increase in yield
efficiency. This latter interpretation is supported by the other observed responses within the
flowering and crop load trial as well as the tipping and fruiting sites trial.
First, in the flowering and crop load trial the number of fruit set per inflorescence
decreased as inflorescence density increased (Figure 4), indicating that the resources
available to each inflorescence decreased as inflorescence density increased.

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 Second, as yield efficiency increased, mean fruit size in the flowering and crop load
trial decreased (Figure 3), indicating that as fruit number increased, the resources available
for each fruit decreased. Crop load effects on fruit size are common across numerous tree
crops, including apple (Breen et al., 2015).
Third, both trials provided evidence of competition for resources between vegetative
and reproductive efforts. In the flowering and crop load trial, as yield efficiency decreased,
canopy volume measured at harvest increased (Figure 5), indicating that as the demand for
resources by fruit production decreased, the resources available for vegetative growth
increased. Results in Year 2 of the tipping and fruiting sites trial showed better spring fruit
set where vegetative shoots were removed, supporting the hypothesis that fruit set in spring
is affected by competition between fruit and vegetative growth, suggesting carbohydrates
are limited at this time. The lack of treatment effects in Year 1, which was characterized by
extensive vegetative growth, may indicate that this competition is only a limiting factor in an
‘on’ year, when overall resource demand from fruit is greater. Alternatively, as trees were
younger and smaller in Year 1, their canopies may have been better illuminated and more
efficient.

Figure 5. The effect of yield efficiency on canopy volume at harvest in the flowering and
crop load trial 2013/14.

In this context, the large difference in the yield efficiencies between the two trials is of
interest. The trees in the flowering and crop load trial were younger and smaller than the
trees in the tipping trial. As avocado fruiting is largely on the periphery of the canopy, the
lower yield efficiency of the older, larger trees may be due to a lower ratio of surface area to
volume of the canopy, as well as increased within-canopy shading and higher maintenance
costs for tree structure. Different seasonal conditions and rootstocks may also have
contributed to the difference.
Crop losses over summer in the tipping and fruiting sites trial made advantages gained
by reducing spring competition between fruit and vegetative shoots short-lived. This result
is consistent with studies by Wolstenholme et al. (1990) and Cutting and Bower (1990). The
retention of some spring-grown leaves in the partial tipping treatments appears not to have
provided sufficient additional resources to ameliorate fruit drop. There were variations
between treatments in the rate of summer fruit drop but not in final fruit retention per shoot
or in yield efficiency (Table 1), suggesting that final fruit numbers are regulated by an overall
resource limitation. Whether summer vegetative growth also competes with fruit was not
addressed in our study but we note that the treatment with the greatest vegetative growth in
Year 2 (T6) also had the highest summer fruit drop (Table 1). T5 also had high vegetative
growth, but low fruit drop over summer, possibly explained by low mean spring fruit set in
this treatment.
Summer drop in ‘Hass’ is significant: the average percentage drop of 38% in Year 1 and

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52% in Year 2 in the tipping and fruiting sites trial is in line with international experience,
recorded in the range 35 to 66% (Wolstenholme et al., 1990; Lahav and Zamet, 1999;
Salazar-Garcia et al., 2006). Understanding and reducing summer fruit drop would appear to
be a key aspect of increasing productivity in avocado.

Flowering sites
The results of the flowering and crop load trial suggest that a strategy for increasing
crop load may be to increase the number of inflorescences by increasing flowering/fruiting
sites. Our attempts to increase fruiting sites in the tipping and fruiting sites trial with the
application of Cytolin to the whole tree were not successful, in contrast to the response to
applications to individual shoots reported by Thorp and Sedgley (1993). This could be due
to practical issues of timing or to the lower rates of application. Alternately, the response to
exogenously applied hormones at the whole tree level may have been limited by
carbohydrate availability for growth at the end of spring.
In addition, in our study there did not appear to be a direct relationship between the
artificial creation of new fruiting sites and increased crop load, suggesting other factors may
be important, for example, the resource costs of branching or the disruption of hormonal
signals. This area requires further study.

CONCLUSIONS
These experiments indicate that improving avocado crop load requires a detailed
understanding of the interactions between inflorescence density, fruit set and retention and
vegetative growth. There appears to be plasticity in the allocation of resources between
vegetative and reproductive effort within the same cropping season. There are also effects
across seasons through the impact of heavy crop loads in both inhibiting return flowering
and reducing the availability of fruiting sites through reduced vegetative growth. The
relative contributions of current and stored carbohydrates in these interactions is not well
understood. Overall, our trials suggest that both flowering intensity and resource limitation
play an important role in determining final crop loads.

ACKNOWLEDGEMENTS
The Small Tree-High Productivity Initiative is an initiative of the Queensland
Government. Major partners include the Department of Agriculture and Fisheries (DAF),
DAF’s research alliance with the University of Queensland (Queensland Alliance for
Agriculture and Food Innovation), and the NSW Department of Primary Industries. A key
element of this initiative has been co-funded by Horticulture Innovation Australia Limited
(Hort Innovation) using the across horticulture levy and voluntary contributions from DAF
and matching funds from the Australian Government through the Hort Innovation project
“Transforming tropical/subtropical tree crop productivity”.
We thank the Donovan Family and Simpson Farms for the use of their trees for trials,
and for their assistance in management and harvest of the crops.

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