GROWING AVOCADOS: FLOWERING, POLLINATION AND FRUIT

SET
Avocados have an unusual flowering behaviour that is affected by
temperature, particularly cold conditions which can impact on the level of
fruit set.
The following article explains the impact of temperature and the use of
pollinisers and pollinators on fruit set of avocados.
FLOWERING AND POLLINATION
A mature avocado tree may produce in excess of a million flowers during
the flowering period, most of which fall without producing fruit. The
purpose behind the mass flowering is to encourage visits by pollen vectors.
In the wild this means a range of flying and crawling insects.
The avocado has a ‘complete’ flower, but with an unusual behaviour
known as ‘protogynous dichogamy’. The avocado flower has both
functional male and female organs in the one flower, but opens and closes
twice over a two-day period — the first day as functionally female (Figure
1) and the next as functionally male (see Figure 2). Each opening stage only
lasts about half a day.
In general, on a single tree all the open flowers will be synchronised. That
means they will be all functionally male or all functionally female. The
avocado pollen of one tree is compatible with itself and quite capable of
pollinating its own flowers — known as self-pollination. The unusual
flowering behaviour is to reduce the likelihood of this occurring — by
minimising the amount of own-pollen about when female stages are
receptive.
To further maximise the likelihood of cross-pollination some trees will
open first in the morning as functionally female, close and then reopen the
next afternoon as functionally male (type A flowering sequence).
Other trees open first in the afternoon as functionally female, close and
then reopen the next morning as functionally male (type B flowering
sequence).
The timing of these stages determines the classification of varieties into
either type A or B flowering. See Table 1 for a diagram of the opening
sequence of type A and B flowering varieties and how this sequence
promotes cross-pollination. This is an evolutionary development to
increase genetic diversity.
It is important to understand that as a ‘clonally’ produced tree, a stand of a
single variety such as Hass will flower essentially as a single tree. That is,
all trees will display the same flowering type behaviour. Table 2 lists some
of the common varieties and their flowering types.
CROSS-POLLINATION FOR GOOD FRUIT SET?
In the normal sequence of events, the avocado tree will produce large
numbers of flowers with some beginning their opening sequence every
day. Because a single flower takes two days to complete its cycle, on any
one day some flowers will be opening as female and others as male, just at
different times of the day.
The stigma of an avocado flower will normally have ceased to be receptive
to pollen when it starts to release its own pollen. However, as a result of
the mass flowering, you will notice from Figure 3 that under ideal
temperatures (maximum 25°C, minimum 20°C) there is theoretically some
overlap from other flowers providing a small window of opportunity for
what is termed close-pollination. Figure 4 provides real-life example of the
cross-over event.
Based on this, many avocado orchards have been planted with Hass only.
However, recent research into increasing avocado production per hectare
has been highlighting the benefits of increasing the period of potential
pollination by inter-planting with complementary varieties, for example, a
type B flowering variety in your Hass orchard.
This is particularly the case if you are in colder locations, or using
management practices that compress the period of flowering, such as the
use of plant growth regulators or ‘scoring’.
This is not a case of just selecting a variety with a known complementary
flowering type to your main variety. Different varieties not only have
either type A or B flowering behaviour, they can also flower at slightly
different periods during the year.
While the opening and closing of flowers over the two-day cycle is
synchronised, it is also affected by air temperature. Under stable ‘ideal’
flowering temperatures (maximum 25°C, minimum 20°C), the flowers
behave much like the typical pattern shown in Figure 3. But Under cooler
conditions (maximum 20°C, minimum 10°C), flower opening can be both
delayed and extended, as shown in Figure 5.
Under even colder conditions, type B flowers may be delayed so much that
the female stage may not be detected, with the flower appearing to open
only as a functional male.
Also, the male phase of type A flowers may be so delayed that they remain
open over-night and into the next morning.
The temperature effect on flower opening is not necessarily equal on all
varieties. Temperatures in the South-West are frequently well below ideal.
As a result of differences for each variety and the importance of achieving
complementary flowering between your main variety (most commonly
Hass) and chosen polliniser, it is beneficial to monitor the flowering
patterns of a range of varieties to determine the best fit under your
conditions.
IDENTIFYING BETTER POLLINISER VARIETIES
In 2009 the flowering of a range of type B avocado varieties was monitored
in the South-West of Western Australia to find how closely they flower in
relation to Hass (Table 3).
This information was based on a single year of observations. While
flowering over the same timeframe does not guarantee they are good
pollinisers because they need to open in synergy under prevailing climatic
conditions, it is a reasonable starting guide.
POLLINISER SPACING
For pollinisers to have the maximum effect they should be of similar
numbers to the complementary variety and evenly spaced within the
orchard. However, often the polliniser variety is not preferred for markets,
or in cooler locations might not yield well if they are type B.
As such, you do not want large numbers of trees producing unpopular or
very little fruit as this is inefficient use of land, water, resources and labour.
If you cannot identify a suitable polliniser that produces fruit in demand
and of acceptable quantities, then aim to plant as few as possible to achieve
acceptable cross-pollination.
The planting pattern of the polliniser variety depends on how the pollen is
expected to move from the polliniser tree to the main variety.
A polliniser tree is located at every third space in every third row (Figure
6). An alternative pattern is a one in eight planting ratio, particularly for
hedge row planting to provide line of sight to a polliniser for all walls of
foliage, so pollinator bees do not have to move through a wall of foliage. A
polliniser tree is located at every fourth space in every second row (Figure
7).
DO WE NEED POLLINATORS?
Yes. The separation in time of the male and female phases of a single
flower and the nature of avocado pollen has led most observers to believe
that a pollen vector or ‘pollinator’ is needed to move pollen from one
flower to another.
The avocado flower provides both pollen and nectar which will attract the
bees, but due to its size, unusual opening sequence and lower sugar
concentration in its nectar, it is not preferred by honey bees. In fact the
ancestral pollinators of avocados were small stingless wasps and flies.
Therefore, while resident populations of feral bees will visit your avocado
trees and do pollination work, they will often be attracted more to other
plants flowering at the same time, thus limiting their effectiveness.
Bees are known to scout an area and learn where preferred plants are and
target these. New bees to an area will not have this knowledge and will
target the closest flowers initially, gradually building their knowledge. To
maximise pollination of your avocados, it is preferable to bring in fresh
hives at the start of flowering (ideally once roughly 10% of flowers have
opened). Do not bring in hives too early as the bees will find other flowers
to visit (eg. Capeweed, Wild Radish, other wildflowers), and even when
the avocado trees begin to flower in earnest, the bees will remain
preferential to the first flowers they found when they were moved into the
area.
The introduction of bee hives into an orchard for pollination requires some
planning. It is preferable to bring in healthy and active hives that are going
to be collecting both pollen and nectar. This is usually done through the
services of a professional pollination provider, who will condition hives
and transport them to your orchard at an agreed time.
As conditioning hives takes time, and flowering can go from 10 to 80%
opened in less than a week, you need to plan in advance with your
pollination provider.
The placement of the bee hives within the orchard also needs to be
considered. They are best placed in small groupings relatively evenly
scattered within the blocks flowering, at a rate of two to five hives per
hectare. Do not place the hives on the outside of the blocks as the scout
bees might find alternative flowers faster and ignore your avocado trees.
Bees are also living insects, so absolute care needs to be taken about the
choice and use of any chemical that may be applied to the orchard during
flowering. If required, always choose the chemical with the lowest bee
toxicity and apply in the evening when bees are less active.
If a bee-toxic chemical is required, you should liaise with the pollination
provider about removing the bees for a few days. 
FRUIT SET
The avocado tree is noted for having large numbers of flowers in some
seasons and yet setting very light crops. There is a range of factors that
affect fruit set. Initially there is the requirement for pollen to be transferred
onto the stigma (female parts) of the flower while it is receptive, as
discussed earlier.
Research has found that increasing the number of pollen grains deposited
onto a receptive stigma will increase the likelihood of effective pollination
— hence the benefit of increasing the number of pollinators and or
pollinisers.
The number of pollen grains being deposited on the stigma is not the end
of the story though, as temperature and humidity are also involved.
Temperature affects the rate of growth of the pollen tube — the lower the
temperature the slower the growth rate.
Temperature and humidity also affect how long before the stigma shrivels.
The higher the temperature and lower the humidity, the quicker the stigma
will shrivel. If the stigma shrivels before the pollen tube has finished its
growth to the ovules then fertilisation will not occur. Dry windy days with
cold nights can lead to this.
There is some suggestion that two to three consecutive days of minimum
temperatures above 10°C, combined with day temperatures above 16°C,
are required to achieve effective pollination. 
Rain and nutrition can also impact on pollination. Rain during the day will
reduce the activity of bees, affecting pollen collection and transferral. This
is similar for cold daytime temperatures as bees are reportedly much less
active at temperatures below 16°C.
Having adequate levels of boron in the floral organs during flowering is
reported to assist in improving fruit set. If low levels are monitored, then
foliar application of boron before flowering may assist.
Additionally, research has shown that when an ovule is fertilised with
pollen from a different variety the embryo that develops from that
fertilisation has 'hybrid vigour' and therefore that fruit is more likely to
remain on the tree and will be a slightly larger fruit than fruits derived
from pollination from the same variety. While other research in California
has shown that this effect does not occur in their conditions it is important
to keep in mind as another reason to plant polliniser trees within the
orchard.
SOME BASIC POLLINATION TERMS
Pollen: The male fertilising agent of plants
Anther: Part of the male organ of the flower from which the pollen is
released
Stamen: The collective term for the anther and filament (stalk)
Stigma: Part of the female organ of the flower onto which the pollen is
deposited
Ovule: female part of the plant which develops into the seed
Embryo: part of the seed which consists of the precursor tissues for the
plant that germinates from the seed.
Pollination: The transfer of pollen from the anther to the stigma
Cross-pollination: Pollen deposited on the stigma is from a different variety
Close-pollination: Pollen deposited on the stigma is from a different flower
of the same variety
Self-pollination: The pollen deposited on the stigma is from the same
flower
Pollinator: The agent which transfers pollen from the male to the female
floral organ (e.g. honey bee)
Polliniser: A cultivar that donates pollen to another cultivar
Dehiscence: The stage when pollen is released from the anthers
Effective pollination: Pollination which leads to fertilisation
Non-effective pollination: Pollination which does not lead to fertilisation
Fertilisation: The fusion of the male gamete with the female gamete
forming the zygote (~seed).