Horticultural Science

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Aspects of avocado fruit growth and development : towards understanding the 'Hass' small fruit syndrome.
(1997) Moore-Gordon, Clive Scott.; Cowan, Ashton Keith.; Wolstenholme, B. Nigel.
Persea americana Mill. cv. Hass is predisposed towards producing a high proportion of undersized fruit. Reasons for phenotypically small 'Hass' fruit are obscure, but it does appear to be aggravated by adverse growing conditions. A detailed study of the metabolic control of avocado fruit growth was carried out to determine the underlying physiological reasons for the appearance of the 'Hass' small fruit phenotype. Furthermore, the application of a mulch was evaluated as a possible management strategy to increase 'Hass' fruit size. Anatomical and morphological comparisons were made between normal and small 'Hass' fruit in an attempt to characterise the 'Hass' small fruit phenotype. Small fruit always contained a degenerate seed coat and fruit size was closely correlated with seed size. Kinetic analysis of changes in cell number and size during fruit development revealed that growth was limited by cell number in phenotypically small fruit. Analysis of endogenous isopentenyladenine (iP) and abscisic acid (ABA) revealed that ABA concentration was negatively correlated with size of similarly aged fruit. Calculation of the iP:ABA ratio showed a linear relationship with increasing fruit size. Qualitative and quantitative differences in mesocarp sterol composition were observed between normal and phenotypically small fruit. Both the normal and small-fruit phenotypes were used to probe the interaction between end-products of isoprenoid biosynthesis and activity of mesocarp 3-hydroxy-3- methylglutaryl coenzyme A reductase (HMGR) in the metabolic control of avocado fruit growth. In phenotypically small fruit, a 70% reduction in microsomal HMGR activity was associated with a substantial rise in mesocarp ABA concentration at all stages of development. Application of mevastatin, a competitive inhibitor of HMGR, via the pedicel reduced growth of phenotypically normal fruit and increased mesocarp ABA concentration. These effects were reversed by co-treatment of fruit with either mevalonate, iP or the synthetic cytokinin (CK) analogue, N-(2-chloro-4-pyridyl)-N-phenylurea, but were unaffected by gibberellic acid. Likewise, in vivo application of ABA reduced fruit growth and HMGR activity, and accelerated abscission at all stages of development, effects that were reversed by co-treatment with iP. In contrast, the effect of sterols on mevastatin-induced inhibition of fruit growth was temporally different. Application of either stigmasterol or cholesterol during phase I caused a decline in growth, accelerated fruit abscission and exacerbated the effects of mevastatin whereas during phase II and III, stigmasterol reversed inhibition of fruit growth. Stigmasterol did not however, reverse the inhibitory effect of mevastatin on HMGR activity - presumably as a result of mevastatin-induced increased endogenous ABA. It was therefore concluded that ABA accumulation downregulates mesocarp HMGR activity and that in situ CK biosynthesis modulates the effect of ABA during phase I of fruit growth whereas, both CK and sterols perform this function during the later stages to sustain the developmental programme. The effect of an altered CK:ABA ratio on solute allocation, cell-to-cell communication and plasmodesmatal structure was investigated in 'Hass' avocado fruits to determine the relationship between a change in hormone balance and expression of phenotypically small fruit. Exogenous application of ABA induced early seed coat senescence and retarded fruit growth, and these effects were negated in fruit co-injected with ABA and iP. The underlying physiological mechanisms associated with ABA-induced retardation of 'Hass' avocado fruit growth included: diminution of mesocarp and seed coat plasmodesmatal branching; gating of mesocarp and seed coat plasmodesmata by deposition of apparently proteinaceous material in the neck region; abolishment of the electrochemical gradient between mesocarp and seed coat parenchyma; and arrest of cell-to-cell chemical communication. In addition, solute allocation in ABA-treated fruit resembled closely that of phenotypically small fruit confirming that elevated ABA concentration had contributed to the decline in postphloem symplastic continuity. In a field trial in the KwaZulu-Natal midlands, root growth was substantially increased throughout three seasons by the application of a coarse composted pinebark mulch. Mulching resulted in a significant 6.6% increase in mean fruit mass, in spite of 14.7% more fruits per tree. The combined effect was a 22.6% increase in overall yield. Differences in productivity between treatments closely correlated to levels of bark carbohydrate reserves. Data collated during this study to suggest that mulching at least partly ameliorated tree stress included: a reduction in the incidence of premature seed coat senescence and pedicel ring-neck, both of which are considered to be advanced symptoms of the stress syndrome; a lowering of mean foliage temperatures; and a reduction in the degree of photoinhibition during the heat of the day.
Ecophysiological studies and tree manipulation for maximisation of yield potential in avocado (Persea americana Mill.)
(1994) Whiley, Anthony W.; Wolstenholme, B. Nigel.; Schaffer, B.
Tree fruit crops generally consist of scion and rootstock components, which through interactive synergism affect tree performance. Coupled with tree architecture, sink/source relationships (both spatial and temporal), genotypic responses to environments, and carry-over seasonal effects present a high level of complexity which often confounds research results. The development, description and use of pheno/physiological models as research and crop management tools is a new holistic approach to reduce complexity and improve understanding of the critical factors which influence crop productivity. A pheno/physiological model is described for cv. Hass avocado growing in a cool, mesic subtropical environment in S.E. Queensland, Australia. Seasonal shoot and root growth had bimodal periodicity with root growth offset and delayed with respect to shoot growth. The priority sink strength of developing shoots compared with roots was confirmed with 14(C) studies. Root growth in summer extended through until late winter when there was a substantial decline following anthesis - a critical time in fruit development with competition between reproductive and vegetative sinks for limited resources. Delayed harvesting of fruit over several seasons resulted in alternate bearing patterns, while removal of fruit at the minimum legal maturity of 21 to 24% dry matter sustained successive high yields. With cv. Hass, production was directly related to starch concentrations in trunks or shoots in July (midwinter) immediately prior to anthesis. However, seasonal starch concentration fluxes in trunks were much lower in coastal subtropical Australia compared with those previously reported from interior areas in more southerly latitudes (7.5% vs. 18% maximum). Current assimilate from over-wintered leaves was necessary to bridge the gap in early spring between the depletion of starch reserves by new reproductive and vegetative shoot growth, and the sink/source transition of the spring shoot growth. Net CO2 assimilation of summer grown leaves reached ca. 17 µmol CO2 m(-2) s(-1), approximately twice as high as previously reported rates on container-grown plants or trees in minimum temperatures were < 10⁰C for 50 days, this being the first report of this phenomenon in field-grown avocado trees. Partial recovery occurred prior to senescence of previous season's leaves in spring after minimum temperatures increased above 10⁰C. The plasticity of the light response was high with the compensation point for net CO2 assimilation at 30 µmol quanta m(-2) s(-1) and the light saturation point at 1270 µmol quanta m(-2) s(-1). Net CO2 fixation from fruit photosynthesis was always less than losses through respiration but was highest during the first few weeks of ontogeny, perhaps contributing to the fruit's own carbon economy at a time when competition for assimilates was greatest. In general, CO2 assimilation studies with current technology applied to orchard trees in non-restrictive soils have elucidated efficiencies more akin to deciduous than evergreen trees - thereby compensating for short-lived leaves and energy expensive fruits. Pheno/physiology models were used to substantiate the most effective timing for trunk injection of ambimobile phosphonate fungicides for the control of Phytophthora root rot, a serious disease of avocados, viz. at the completion of the leaf expansion phases when leaves were strong net exporters. Preliminary studies demonstrated potential yield increases when the assimilation efficiency of photoinhibited over-wintered leaves was improved through increased nitrogen concentration, and spring shoot growth was partially suppressed with foliar sprays of the growth retardant paclobutrazol.
Special carbohydrates of avocado : their function as 'sources of energy' and 'anti-oxidants'.
(2009) Tesfay, Samson Zeray.; Bertling, Isa.
There is increasing interest in special heptose carbohydrates, their multifunctional roles from a plant physiological view point in fruit growth and development as well as in the whole plant in general due to their potential in mitigating photo-oxidative injury to the whole plant system and the image of avocado as ‘health fruit’. Studies have been carried out to investigate the role of avocado heptoses, rare carbohydrates predominantly produced in avocado. Several authors have documented various research findings and speculated on multifunctional roles of avocado special sugars. However, few reports have made an attempt to elucidate the multifunctional roles of avocado heptose carbohydrates as: ‘sources of energy’, storage and phloem-mobile transport sugars, and precursors for formation of antioxidants. Assessing the avocado carbohydrates over the plant growth and development during ontogeny may, therefore, offer clues to better understand whole plant behaviour. Plant sampling was carried out over different developmental stages. Using plants grown in the light versus etiolated seedlings; sugar determinations were also done to determine what sugar is produced from which storage organs. The sugars were extracted and analysed by isocratic HPLC/RID. The embryo had 47.11 % hexose and 52.96 % heptose sugars. The seed, however, also released significant amounts of D-mannoheptulose (7.09 ± 1.44 mg g-1 d. wt) and perseitol (5.36 ± 0.61 mg g-1 d. wt). Similarly fruit and leaf tissues had significant amounts of heptoses relative to hexoses at specific phenological stages. In postharvest ‘readyto-eat’ fruit the following carbohydrate concentrations were as follows:exocarp heptoses 13 ± 0.8; hexoses 4.37 ± 1.6 mg g-1 d. wt, mesocarp heptoses 8 ± 0.2; hexoses 3.55 ± 0.12 mg g-1 d. wt), seed heptoses (only perseitol) 13 ± 1.1; hexoses 5.79 ± 0.53 mg g-1 d. wt. The results of this experiment was the first to demonstrate that the heptoses D-mannoheptulose, and its polyol form, perseitol, are found in all tissues/organs at various phenological stages of avocado growth and development. Secondly, heptoses, as well as starch are carbohydrate reserves that are found in avocado. The heptoses, beyond being abundantly produced in the avocado plant, are also found in phloem and xylem saps as mobile sugars. The study also presents data on the interconversion of the C7 sugars Dmannoheptulose and perseitol. It is deduced that D-mannoheptulose can be reduced to perseitol, and perseitol can also be oxidized to D-mannoheptulose by enzymes present in a protein extract of the mesocarp. The potential catalyzing enzyme is proposed to be an aldolase, as electrophoretic determinations prove the presence of such an enzyme during various stages of development in various plant organs. Avocado heptoses play an important role in plant growth and development and in fruit in particular. Moreover, they are reported as sources of anti-oxidants, and contribute significantly to fruit physiology if they function in coordination with other anti-oxidants in fruit tissues. To evaluate the presence of anti-oxidant systems throughout avocado fruit development, various tissues were analysed for their total and specific anti-oxidant compositions. Total anti-oxidant levels were found to be higher in the exocarp and in seed tissue than in the mesocarp. While seed tissues contained predominantly ascorbic acid (AsA) and total phenolics (TP), the anti-oxidant composition of the mesocarp was characterised by the C7 sugar, D-mannoheptulose. Among the anti-oxidant enzymes assayed, peroxidase (POX) and catalase (CAT) were present in higher concentrations than superoxide dismutase (SOD) in mesocarp tissue. Different anti-oxidant systems seem to be dominant within the various fruit tissues. Carbohydrates are the universal source of carbon for cell metabolism and provide the precursors for the biosynthesis of secondary metabolites, for example via the shikimic acid pathway for phenols. The preharvest free and membrane-bound phenols, catechin and epicatechin, are distributed differently in the various fruit tissues. Membrane-bound and free phenols also play a role as anti-oxidants, with free ones being more important. KSil (potassium silicate) application to fruit as postharvest treatment was used to facilitate the release of conjugates to free phenols via lysis. This treatment improved fruit shelf life. Western blotting also revealed that postharvest Si treatment affects the expression of enzymatic anti-oxidant-catalase (CAT). Overall the thesis results revealed that C7 sugars have anti-oxidant properties and that D-mannoheptulose is the important anti-oxidant in the edible portion of the avocado fruit. Dmannoheptulose is furthermore of paramount importance as a transport sugar. Perseitol on the other hand acts as the storage product of D-mannoheptulose, which can be easily converted into D-mannoheptulose.

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