Proceedings of Third World Avocado Congress 1995 pp. 3-6.
UPTAKE
OF PGRs INTO DETERMINATE VERSUS INDETERMINATE INFLORESCENCES OF THE 'HASS'
AVOCADO (PERSEA AMERICANA MILL.)
I.
Bertling and C.J. Lovatt
Department of Botany and Plant Sciences, University
of California,
Riverside, CA 92521-0124, USA
Additional
index words
Abstract
The development of the
vegetative shoot apex of the indeterminate inflorescence of the avocado
potentially renders transpiration, water flux, and sink-source relationships
within this type of inflorescence different from those of the determinate
inflorescence. The influence of indeterminate inflorescence leaves on the
import and distribution of xylem-transported PGRs within the inflorescence is
not known. Thus, in this study, the uptake of tritiated PGRs into flowers,
setting fruit, and small fruit of 'Hass' avocado (Persea americana Mill.)
was contrasted in the two inflorescence types.
The main axis of intact
determinate and indeterminate inflorescences at advancing stages of development
were placed into small plastic jars containing one of the following tritiated
PGRs: abscisic acid, indole-3-acetic acid, gibberellic acid, isopentenyl
adenosine or zeatin riboside. Transport and allocation to the different organs
of the inflorescences was determined.
At flower opening, greater
amounts of IAA and ABA (on a per g fresh weight basis) were taken up by the
developing leaves of the indeterminate inflorescence, but the amounts allocated
to the flowers of either inflorescence type were not statistically different.
At early fruit set, the vegetative flush continued to have a greater
concentration of ABA than other organs. Indeterminate fruitlets accumulated GA3,
IPA, and ZR at concentrations equal to the leaves, but greater than fruitlets
of determinate inflorescences. At later stages of fruit development, leaves
still had greater concentrations of GA and IPA, but fruit on indeterminate and
determinate inflorescences had equal concentrations of all PGRs.
Taken together, the results
suggest that the vegetative flush does not deprive the flowers borne on
indeterminate inflorescences of PGRs during early fruit development and that
PGRs are allocated to flowers and fruit on the basis of a sink strength rather
than transpirational flux.
1.
Introduction
Interactions between the
different organs of a fruit tree are complex. In avocado, competition between
young developing fruit and the spring flush of vegetative growth has been
implicated in reducing fruit set (Kalmar and Lahav, 1976; Köhne,
1987; Cutting
and Bower, 1989). Conversely, a heavy fruit set reduces the intensity of the
subsequent summer and fall vegetative flushes (Whiley et al., 1988). In
addition, fruit compete for resources among themselves (Wright, 1989). The involvement
of plant growth regulators in these regulatory processes has been long
established (Naylor, 1984).
The two
inflorescence types of avocado (the determinate, totally floral, shoot and the
indeterminate floral shoot terminating in a vegetative apex), provide a useful
system to investigate the effect of the vegetative apex on the transport and
distribution of plant growth regulators within the inflorescence.
Our objective was
to determine quantitatively if the partitioning of PGRs supplied via the
vascular system to flowers and fruit of an inflorescence is influenced by the
presence of leaves, and if this partitioning changes during development.
2. Material and methods
Adult avocado trees of the
cultivar 'Hass' located at the University of California, Riverside, were
selected for the experiment. Determinate and indeterminate inflorescences were
collected from trees and placed into plastic receptacles containing 6 ml of a 0.01
M 2-(N-Morpholino)ethanesulfonic acid (MES) solution, pH 6.4. To this solution,
2x105 dpm of one of the following tritiated plant growth regulators was added:
indole-3-acetic acid (IAA), abscisic acid (ABA), gibberellic acid (GA3), zeatin
riboside (ZR), and isopentenyl adenosine (IPA). Each PGR was previously
demonstrated to occur in the vascular system (Bertling and Lovatt, in prep.)
The experiment was carried
out at three different stages of development: (i), at full bloom when most of
the flowers were open and the vegetative flush had just started to grow; (ii),
at petal fall when the first signs of fruit set were visible and the
inflorescence leaves were rapidly expanding; and (iii), when fruit were
2 by 3 cm (w x 1) in size and the oldest leaf was fully expanded.
After an incubation period of
24 h under constant light at 500 μmol/mm2/s and 30°C,
flowers/fruit and leaves were detached from the inflorescence and immediately
shock frozen. Samples were homogenized, extracted in methanol over night, and
thereafter the methanol removed in a rotary evaporator. The residue was
dissolved in 0. 1 M acetic acid, centrifuged at 15x 103 rpm for 15 min. The
supernatant fraction was further purified through a C18 SepPak cartridge and
the filtrate dried in a vacuum concentrator. The dried extract was dissolved in
methanol, diluted with liquid scintillant, and the content of radioisotope
measured with a liquid scintillation spectrophotometer.
3. Results
The uptake and translocation
of GA3 into flowers and fruit was greater than any other PGR at all
stages of inflorescence development with the single exception of the
concentration of IAA observed for fruit at stage III (Table 1). At stage I, GA3
concentrations in flowers in either inflorescence type were not significantly different.
After petal fall (stage II), indeterminate fruitlets contained GA3
concentrations equal to those of the leaves and significantly greater than
fruit borne on determinate inflorescences. At stage III, GA3 concentrations
were the same in fruit borne on both determinate and indeterminate
inflorescences, but significantly greater in the leaves despite the fact that
they were approaching full expansion.
Through all stages of
inflorescence development, leaves of indeterminate inflorescences exhibited significantly
greater concentrations of ABA than flowers or fruit of either inflorescence
type (Table 1). The accumulation of ABA by leaves, which decreased from stage I
to III as the leaves expanded, had no impact on the uptake, translocation or
distribution of ABA into flowers or fruit borne on the two types of
inflorescences, i.e., their ABA concentrations were not statistically different
at any stage of development.
Transport of ZR into leaves
of indeterminate inflorescences was significantly greater than into flowers and
fruit of these inflorescences at stages I and III. ZR was the only PGR that
accumulated in flowers and fruit of determinate inflorescences to a
concentration equaling that of the leaves of indeterminate inflorescences
(stages I and III). Relative to ZR, the uptake and translocation of the
cytokinin IPA was consistently lower. At stage II, fruit and leaves of
indeterminate inflorescences had similar concentrations of ZR and IPA,
respectively. These ZR and IPA concentrations were significantly greater than
those of fruit borne on determinate inflorescences. By stage III, fruit borne on
indeterminate inflorescences had ZR and IPA concentrations equal to fruit of
determinate inflorescences and significantly lower than their leaves.
IAA uptake and
translocation into leaves was initially high (stage I) and greater than in the
flowers of either inflorescence type. At later stages of development, no
significant differences between organs could be detected.
By stage III, PGR
concentrations of fruit home on determinate versus indeterminate inflorescences
were not significantly different. Consistent with this, the number of fruit
retained by each inflorescence type was not significantly different: compare
3.9+/-1.5 to 2.6+/-0.9 fruitlets per determinate and indeterminate
inflorescences, respectively.
4.
Conclusions
In general, uptake and
translocation of the PGRs analyzed in this study was greater into leaves of the
vegetative apex of the indeterminate inflorescences than into flowers or fruit borne
on either inflorescence type over the various stages of development. An unusual
but consistent exception was the concurrent accumulation of PGRs (GA3, ZR, and IPA) in fruit of
indeterminate inflorescences at stage II of development. These differences
disappear by stage III, resulting in leaf concentrations of GA3, ZR, and IPA
that were greater than those determined for the indeterminate fruit. The
relatively high leaf concentrations of GA3, ZR, IPA, and ABA at stage III are
of interest as the leaves were approaching full expansion and maturation,
consistent with the well known movement of these PGRs in the transpiration
stream. Worthy of note is the fact that IAA, which is not known to move by
transpirational flux, was somewhat lower in the leaves of indeterminate
inflorescences at stage III.
The greater translocation of
PGRs into indeterminate inflorescences due to the presence of leaves, resulted
in greater PGR concentrations in the fruit borne on indeterminate
inflorescences in several cases cited above. Reduced levels of PGRs in these
fruit were not observed at any stage. This result is consistent with the fact
that reproductive organs are greater sinks than vegetative organs (Cannell,
1985).
Acknowledgements
This research was supported
in part by a grant from the California Avocado Development Organization, the
California Avocado Commission, the Citrus Research Center and the Agricultural
Experiment Station of the University of California, Riverside. The authors
thank Dr. Jonathan Cutting for generously donating several 31-1-
PGRs used in this study.
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Table
I - Uptake, translocation and distribution of 3H-PGRs in determinate
(D) and indeterminate (I) inflorescences of 'Hass' avocado from flowering to fruit
set. Data are presented in dpm per g flowers, fruit or leaves per
inflorescence, and are the average from 6 separate experiments of 6
inflorescences, at 3 different dates.