South African Avocado Growers’ Association Yearbook 1987.
10:42-43.
Proceedings of the First World Avocado Congress
M SEDGLEY
Department
of Plant Physiology, Waite Agricultural Research Institute,
Glen Osmond,
SA, 5064, Australia
Research has shown that all stages of the reproductive cycle of avocado
are temperature sensitive. The tree appears to perform best under a temperature
regime of approximately 25°C day and 20°C night. This knowledge of the limits
of sensitivity of the various stages of floral initiation, floral development,
pollen tube-growth and fruit-set present the opportunity to select the correct
cultivar for the environment. In addition, the information can be used by plant
breeders as selection criteria.
Yield of avocados is dependent upon successful floral initiation, floral development, pollination and fruit-set. Problems with any of these processes will have a detrimental effect on fruit production which cannot be alleviated by alternative cultural practices. It is important to understand the physiology of the reproductive processes so that the management of the crop includes provision for optimising yield.
Successful floral initiation and
development are essential processes in the reproductive cycle. In the
Mediterranean climate of southern Australia floral initiation in the cultivar
Fuerte occurred in autumn during April and May and flowering occurred in spring
during September to November (Scholefield, Sedgley & Alexander, 1985).
Vegetative buds had a shallow
terminal meristem with vegetative axillary meristems. During the transition to
flowering in April and May elongation of the secondary axillary meristems
occurred and tertiary floral axes were produced. Differentiation of the first
individual flower buds at the tips of the secondary and tertiary axes was
observed in May. The terminal meristem of the bud remained vegetative and only
the axillary meristems underwent the transition to flowering. During anthesis
in spring the floral axillary meristems were the first to burst, followed by
the terminal vegetative meristem of the bud which contributed to the continued
extension growth of the tree.
There were two main vegetative
growth flushes per year, one in summer just prior to floral initiation and one
in spring coinciding with the end of flowering. Carbohydrate levels in the
trunk and branches were lowest just prior to floral initiation. Carbohydrates
accumulated during the period of floral development in winter, reached maximum
levels just prior to flowering and decreased again during the fruit development
period. The biennial bearing habit appeared to be related to the carbohydrate
status of the tree as heavy yields followed high levels of starch accumulation
and low yields followed low levels.
Floral initiation and development
is not a problem in southern Australia but poor flowering is regularly observed
in the tropical climate of northern Australia (Sedgley, Scholefield &
Alexander, 1985). Terminal buds sampled during spring were floral and contained
flowers which had differentiated perianth primordia. Thus floral initiation was
occurring but the floral buds did not burst. Vegetative extention growth of the
tree occurred not from the terminal vegetative meristem of the floral bud but
from lateral buds lower down the shoot. This abnormal growth pattern gave the
trees a rather straggly appearance. The problem was particularly marked amongst
Mexican and Guatemalan type cultivars; less so in West Indian types.
To test the hypothesis that the
inhibited reproductive development was caused by high temperature, avocado
plants were grown under controlled environmental conditions. Under a tropical
temperature regime of 33°C day and 23°C night the trees of the cultivars Fuerte
and Hass had fewer flowers and a shorter flowering period than under
temperature conditions of 25°C day and 15°C night. The features of unburst
floral buds and lateral vegetative extension growth observed in the orchards in
northern Australia were reproduced under the controlled environmental
conditions. The unburst floral buds were inhibited at the stage of stamen
differentiation and it was concluded that in the avocado this stage was
particularly sensitive to high temperature.
Avocado flowers show protogynous
dichogamy with the female stage followed by a closed stage followed by the male
stage. There are two flowering types; type A, with the female stage in the
morning, and type B with the female stage in the afternoon. Interplanting of
type A and type B cultivars in the orchard ensures that cross-pollination will
occur. The floral cycle of the avocado is very sensitive to temperature
fluctuations. In the Fuerte cultivar daytime temperatures above 30°C or below
20°C were found to disrupt flowering (Sedgley, 1977a). High temperatures
appeared to stimulate vegetative growth at the expense of reproductive
development and flowers and developing fruit were shed from the plant. Low
temperatures were particularly damaging as less than 10 per cent of the flowers
opened in the female stage. Other type B cultivars including Ryan, Edranol,
Sharwil and Hazzard also showed female sterility at a daytime temperature of
17°C, although the cultivar Bacon was an exception and had a high proportion of
female stage flowers under these conditions (Sedgley & Grant, 1983).
Type A cultivars, including Hass,
Reed, Wurtz, Rincon & Jalna responded better than the type B cultivars to
low temperatures during flowering (Sedgley & Annells, 1981; Sedgley &
Grant, 1983). All had female stage flowers although the floral cycle was
extended to over double the normal length in some cases and the main periods of
anthesis occurred during the night. This would reduce pollination and fruit-set
as pollinating insects would not be active during the night.
Successful fertilisation and
fruit-set results from pollination of the female stage flower; the male stage
flower is not female fertile (Sedgley, 1977b). The optimum temperature for
pollen tube growth of all cultivars tested was 25°C (Sedgley, 1977a; Sedgley
& Annells, 1981; Sedgley & Grant, 1983). Pollen tube growth to the base
of the style was rapid and occurred within three hours, but penetration of the
ovule by a pollen tube did not occur until 18 to 24 hours after pollination
(Sedgley, 1979b). All cultivars tested were cross-compatible and no
pollen-pistil incompatibility was detected between any cultivars (Sedgley,
1979a). There were differences in female fertility, however, as the cultivar
Fuerte had a higher proportion of defective embryo sacs than Hass and this
resulted in a lower proportion of fertilised fruits following pollination
(Sedgley, 1979b).
A major problem in avocado
production is the high rate of post-pollination fruit shed. The majority of
fruits are shed during the month following anthesis although some fruit
abscission can occur throughout the developmental period of fruit growth. The
anatomy of abscissed fruits was compared with that of actively growing fruits
to investigate any possible reasons for the high rate of fruit shed (Sedgley,
1980). During the first weeks after anthesis the majority of the flowers were
unfertilised. Less than 20 per cent of the flowers were abnormal and poor
pollen transfer was identified as the major reason for this early drop.
However, by one month following anthesis all shed fruit were fertilised and had
normal embryo and endosperm development. Some of the shed fruits showed some
degeneration but this probably occurred during the period between the cessation
of growth of the fruit and abscission from the plant. This period was
approximately one week. The majority of the shed fruit had reached a stage of
embryo and endosperm development corresponding to 14 days after pollination. No
anatomical reason for the high rate of fruit shed was observed and it is suggested
that competition effects may be responsible for abscission. There is
competition not only between developing fruits, but also between the fruits and
vegetative flush growth. Following the bursting of the floral axillaries of the
buds, the vegetative terminal bursts to continue the extension growth of the
shoot. This bud develops expanding leaves which are in direct competition with
the young fruits and also enjoy a position apical to the fruits. It is possible
that this competition situation may be at least partly responsible for the high
rate of early fruit shed.
This work on the reproductive
biology of the avocado has benefited the avocado industry in Australia and
overseas. In particular the work on temperature sensitivity of flowering in
avocado has identified the importance of selecting the correct cultivar for the
environment. The work on temperature at flowering time has been further
developed for the Australian situation and is valuable information for the
Australian industry (Whiley & Saranah, 1986). In addition to the
application in improving orchard yields, the work also has relevance to the
plant breeding situation. Cultivars and breeding lines can be selected on the
basis of their temperature sensitivity during reproductive development and so
selection of cultivars for certain areas can be refined. In addition a
knowledge and understanding of the breeding system of a crop enables greater
manipulation of the species for the improvement of plant breeding.
1 Scholefield, PB, Sedgley, M
& Alexander, D, McE, 1985. Carbohydrate cycling in relation to shoot
growth, floral initiation and development and yield in the avocado. Scientia Horticulturae, 25, 99-110.
2 Sedgley, M, 1977a. The effect of
temperature on floral behaviour, pollen tube growth and fruit set in the
avocado. Journal of Horticultural Science,
52, 135-141.
3 Sedgley, M, 1977b. Reduced
pollen tube growth and the presence of callose in the pistil of the male floral
stage of the avocado. Scientia
Horticulturae, 7, 27-36.
4 Sedgley, M, 1979a.
Inter-varietal pollen tube growth and ovule penetration in the avocado. Euphytica, 28, 25-35.
5 Sedgley, M, 1979b. Light
microscope study of pollen tube growth, fertilisation, and early embryo and
endosperm development in the avocado varieties Fuerte and Hass. Annals of Botany, 44, 353-359.
6 Sedgley, M, 1980. Anatomical
investigation of abscissed avocado flowers and fruit lets. Annals of Botany, 46, 771-777.
7 Sedgley, M & Annells, CM,
1981. Flowering and fruit-set response to temperature in the avocado cultivar
Hass. Scientia Horticulturae, 14, 27-33.
8 Sedgley, M & Grant, WJR,
1983. Effect of low temperatures during flowering on floral cycle and pollen
tube growth in nine avocado cultivars. Scientia
Horticulturae, 18, 207-213.
9 Sedgley, M, Scholefield, PB
& Alexander, D McE, 1985. Inhibition of flowering of Mexican and
Guatemalan-type avocados under tropical conditions. Scientia Horticulturae, 25, 21-30.
10 Whiley, AW & Saranah, JB,
1986. Varietal performance and future developments with new varieties. Maroochy Horticultural Research Station Report, No 4, 30-37.
Queensland Department of Primary Industries.