Doctoral Degrees (Horticultural Science)

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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.
Management of avocado postharvest physiology.
(2011) Blakey, Robert John.; Bower, John Patrick.; Bertling, Isa.
Avocados are an important horticultural crop in South Africa, especially in the provinces of KwaZulu-Natal, Mpumalanga and Limpopo. The distance to traditional export markets, phytosanitary restrictions to lucrative markets such as China, the USA and Japan and increased competition in the European market have challenged the South African avocado industry. The industry has responded with improved logistics and shipping, a co-ordinated market access program and a global system to co-ordinate exports of avocados to the European market. To remain competitive on the global market, further improvements and innovations are required to improve the efficiency of postharvest operations. These improvements and innovations should be guided by a greater understanding of postharvest physiology. Avocados are a relatively new export crop, so there is still much to be learnt about avocado postharvest physiology and the optimisation of postharvest management. In this regard, reduced temperature storage (1°C) and modified humidity packaging (MHP) were investigated for their effect on fruit physiology and quality, the effect of a water- and ABA-infusion on ripening was examined and the effect of a cold chain break on fruit physiology and quality determined; near-infrared spectroscopy was also examined for its potential for its use in the avocado industry. As an initial study, the relationships between individual sugars, protein and oil were studied to understand the changes in avocado fruit during ripening. It was found that mannoheptulose and perseitol were the predominant sugars at harvest, but declined to very low levels during the first 10 days postharvest. The concentrations of glucose and fructose increased, while sucrose declined slightly during ripening. The concentration of protein increased sigmoidally during ripening, reflecting the increase in the ripening enzymes, particularly cellulase and polygalacturonase. The oil content fluctuated slightly during ripening. It is suggested that mannoheptulose and perseitol are important carbon and energy sources during ripening. Glucose concentration was also found to increase earlier in fast ripening fruit compared to slow ripening fruit, which is related to increased cellulase activity and may be related to the ABA functioning. Thereafter, storage and ripening trials in two consecutive seasons showed that 1°C storage and the use of MHP for 28 days reduced mass loss, water loss, ethylene production, respiration, softening and heptose consumption, without appreciably affecting fungal rots, physiological disorders or external chilling injury, compared to fruit stored at 5.5°C and regular atmosphere respectively. Also, the storage of fruit in MHP delayed the rise in the activity of cellulase during ripening, compared to fruit not stored in MHP, but there was no significant difference in the peak activity of cellulase, polygalacturonase or pectin methylesterase. In a separate experiment, fruit ripening was significantly affected by the infusion of ABA in an aqueous solution. Water slightly reduced the variation in ripening while ABA reduced the time to ripening and the variation; it is suggested that water stress and ABA are intrinsically involved in the ripening processes and may act as a ripening trigger. The water concentration in fruit was measured non-destructively using reflectance NIR; this model was used to determine the maturity of fruit and the loss of water during cold storage. In the cold chain break experiment, it was found that although fruit recovered after a cold chain break, in terms of ethylene production and respiration, there was a loss in quality because of severe shrivelling as a result of increased water loss. Fruit that were stored at 1°C were generally of a better quality at ripeness, if the cold chain was broken, compared to fruit stored at 5.5°C. In a follow-up experiment, it was found that significant changes occurred in avocado physiology over a 6h period. The respiration rate of fruit significantly increased after 4h at room temperature and mannoheptulose declined by 32% in control fruit and by 16% in ethephon-treated fruit after 6h. This demonstrates the potential for quality loss in a short amount of time. Furthermore, a model of avocado ripening is proposed, outlining the role of water, ABA, ethylene, respiration, ripening enzymes and individual sugars. This study has contributed to the understanding of avocado postharvest physiology and should aid in better management of avocados for improved fruit quality and consumer satisfaction.
Optimization, validation and application of radioimmunoassays for plant growth substances in avocado (Persea americana Mill.) fruits.
(1984) Cutting, Jonathan Garth Melville.; Wolstenholme, B. Nigel.; Lishman, Arthur William.
The objective was to develop, optimize and validate radioimmunoassays (RIA) for several plant growth substances (PGS) and then apply the RIA's to determine PGS trends in 'Fuerte' avocado fruits from fruit set to fruit ripening. Antibodies to the cytokinin isopentenyl adenosine (IPA) were obtained from rabbits inoculated with a periodate-derived IPA-BSA conjugate. The antiserum cross-reacted (25%) with only 2iP (isopentenyl adenine). The RIA measuring range was from 0,1 to 100 ng. Anti-IPA serum was used to develop a RIA for 2iP, with a measuring range from 0,5 to 100 ng. Using Dowex 50W-X8 and cellulose TLC purified avocado fruit extract, 20 samples per day could be processed. The RIA for abscisic acid (ABA) was developed from rabbit antibodies from an inoculated carbodiimide-derived, (+ -) ABA-BSA conjugate. The free active ABA component was isolated prior to quantitation by RIA by solvent partitioning and silica gel TLC. The indole-acetic-acid (IAA) RIA was established from sheep-produced antibodies to a formaldehye-derived IAA-BSA conjugate, after repeated inoculations. For both the ABA and IAA RIA's, contaminants in the avocado tissue were removed by solvent partitioning. Developing avocado fruits, and especially young fruits were rich sources of IAA and 2iP in particular with seed concentrations exceeding those of the fruit flesh. The concentration of ABA rose throughout fruit development, reaching 100 ng g(-1) in the flesh at fruit maturity. Just prior to seed and fruit maturity, relatively high levels of IAA, 2iP and IPA were associated with the thick, fleshy testa, rapidly declining to zero as the testa dried. The avocado fruit is physiologically dependent on the seed right up to this stage, and testa maturity correlates well with minimum "legal" maturity of 80% moisture content of flesh. In ripening avocado fruit, the concentration of free ABA rose as softening progressed. Total ABA concentrations fell initially, and then rose after a firmometer reading between 50 and 60 (100 coincides with eating ripeness). Later harvested fruit had double the ABA concentration of early harvested fruit. The ripening stimulus appeared to be related to moisture stress in the fruit. This study has confirmed the prime advantages of sensitivity, specificity and rapidity of RIA, as well as its usefulness in multi-PGS studies and batch-type analysis.
Plant hormone homeostasis and the control of 'hass' avocado fruit size.
(2002) Taylor, Nicolette Jane.; Cairns, Andrew Lawrence Patrick.; Cowan, Ashton Keith.; Van Staden, Johannes.
The 'Hass' avocado produces two distinct phenotypically different populations of fruit, i.e. normal and small fruit. The small fruit variant is characterized by early seed coat senescence that results in arrested growth, due to dramatically reduced cell cycle activity. This system has been used to study the metabolic control of fruit growth for two reasons. Firstly, the 'Hass' avocado is a major export crop in South Africa and unmarketable small fruit cost the industry millions of rands per season. Secondly, in the absence of evergreen tree-crop mutants with which to dissect controlling mechanisms contributing to the control of final fruit size, the 'Hass' avocado and its small fruit variant provides an ideal system to investigate the physiology, biochemistry and molecular biology of fruit growth in subtropical species. A detailed study was conducted to probe the contribution of hormones in the control of final fruit size by comparing and contrasting tissue distribution and content of hormones in developing 'Hass' avocado and its small fruit variant. In addition the proposal that changes in hormone homeostasis occur as a result of differences in the allocation of the molybdenum cofactor (MoCo) and changes in the activity of xanthine dehydrogenase (XDH) and the aldehyde oxidases (AO) involved in abscisic acid (ABA) and indole-3-acetic acid (IAA) metabolism was evaluated. Activity of XDH, xanthoxal (XAN) oxidase, indole acetaldehyde (IA-aid) oxidase and cytokinin oxidase (CKOX) was related to tissue content and composition of IAA and ABA. Comparisons between normal and small fruit revealed that under conditions where CKOX is elevated, the increased adenine produced inhibits XDH activity, which leads to elevated activity of the AOs involved in ABA and possibly IAA biosynthesis as a result of increased MoCo allocation to these enzymes. Further analyses revealed that both cytokinin (CK) and auxin elevates CKOX activity and that adenine and CK do indeed inhibit XDH activity, which leads to increased AO activity. In addition, application of CK to normal fruit increased IAA in mesocarp tissue but reduced IAA content of seed tissue and reduced ABA in mesocarp tissue but had no effect on ABA in seed tissue. Cytokinin oxidase therefore contributes to the regulation of ABA and IAA metabolism during plant organ growth by modulating the activity of XDH. Low XDH and lA-aid oxidase activity together with high XAN oxidase and CKOX activity early in fruit development combine to reduce both elongation and radial growth, which results in the appearance of the 'Hass' small fruit phenotype. This event was associated with high ABA and low IAA in seed tissue of small fruit, but high ABA and IAA in seed coat and mesocarp tissue of these fruit. Thus, whilst low IAA in seed tissue is associated with reduced growth the . reverse is true in seed coat and mesocarp tissue where high IAA retards tissue growth. Calculation of CK/ABA and CK/IAA ratios revealed that a decrease in these ratios was found in mesocarp tissue of small fruit. However, in seed tissue of small fruit both IAA and ABA were decreased relative to CK. The maintenance of the correct hormonal balance in avocado fruit thus ensures the continuation of cell division cycle activity, with any changes responsible for the high incidence of a small fruit variant in the 'Hass' avocado.
Post-phloem transport and metabolism of sucrose in avocado.
(2001) Cripps, Ryan F.; Smith, Michael Trevor.; Cowan, Ashton Keith.
In South Africa, and in several other sub/tropical countries, the avocado represents a commercially important crop. Very little is currently understood about the metabolism of sugars in this fruit. The variety 'Hass' is a popular cultivar that is grown extensively in South Africa. However this. cultivar has a tendency to produce two distinct fruit phenotypes: a normal sized variant and a small, undersized variant. Current literature suggests that the small fruit phenotype is characterised by an elevated abscisic acid (ABA) to cytokinin ratio and altered isoprenoid metabolism. The results presented in the current investigation represent the findings from a detailed study into the metabolism and transport of sugars in 'Hass' fruit in an attempt to characterise solute allocation in developing avocado fruit. Furthermore, the activities of sugar metabolising enzymes, routes of solute movement and polyphenolic contents of normal, small and ABA-treated fruit were compared and contrasted to evaluate the potential role of ABA in the induction and expression of the small fruit phenotype. The enzymes invertase, sucrose synthase (SSy) and sucrose phosphate synthase are involved in the metabolism of sucrose (Suc) and, hence, phloem unloading, post-sieve element transport and fruit growth. Although not the major sugar present, Suc was found in avocado phloem sap, and the enzymology for its metabolism was shown to exist in avocado fruit. It appears that sink strength is established during early fruit growth by high acid invertase activity, especially during the period of rapid cell division. As fruit growth progresses the activity of SSy and an enzyme responsible for the oxidation of perseitol (tentatively termed perseitol dehydrogenase) increases, suggesting that these enzymes play an important role in the supply of carbon during the linear phase of fruit growth. All Suc metabolising enzyme activity diminishes as the fruit approaches maturity. With the exception of SSy (in the cleavage direction), all enzymes assayed showed a general increase in relative rates of activity in small and ABA-treated fruit. Similarly, ABA-treatment of seed coat discs in vitro resulted in the elevation of insoluble and soluble acid invertase, SSy (in the synthesis direction), and sucrose phosphate synthase activity. Furthermore, both small and ABA-treated fruit were characterised by elevated total soluble sugars, glucose and fructose levels. These observations suggest that altered sugar metabolism, as a consequence of changes in endogenous ABA levels, may contribute to the occurrence of the small fruit. The seed coat represents an import link between the seed, the mesocarp and the parental plant tissues. Loss of seed coat and endosperm integrity accompanied fruit maturation and a reduction in the movement of solutes into the seed. An increase in polyphenolics in the seed coat tissue seemed critical in this reduced movement. Both the small and ABA-treated fruit were characterised by early senescence of the seed coat, which was accompanied by both a loss of transport into and out of the seed and premature maturation of the fruit. This premature seed coat senescence appeared similar to programmed cell death in tissues exposed to stress or elevated reactive oxygen species, stimuli that are often accompanied by elevated ABA levels. Callose was localised to the plasmodesmata and is proposed to play a role in the gating of, and hence movement through, these pores. Small fruit were characterised by a loss of symplastic continuity, as represented by fewer plasmodesmata, and reduced callose degradation. Comparison of callose content and rates of synthesis suggest that ABA-treatment, similarly, reduces callose catabolism. The association of ABA with both the premature senescence of the seed coat and a reduction in symplastic continuity, and, hence, a reduction in solute transport, further cements the potential role of ABA in the occurrence of the small fruit phenotype.
Some aspects of water relations on avocado Persea americana (Mill.) tree and fruit physiology.
(1985) Bower, John Patrick.; Wolstenholme, B. Nigel.
The effect of long-term irrigation on water relations of 'Fuerte' avocado trees, and the consequences for fruit ripening and physiology, particularly physiological disorders, were studied. Four irrigation regimes were used, namely dryland relying on rainfall, occasional (irrigation when soil moisture tension reached 80 kPa), regular (soil moisture tension 55 kPa) and frequent (soil moisture 35 kPa). Seasonal tree leaf water potential was studied. This became more negative during the dry season (winter and spring) and less negative during the period of summer rainfall. During the dry periods, the dryland and 80 kPa treatments had considerably more negative leaf water potential than the 55 kPa and 35 kPa regimes. These differences decreased during the summer rainfall period. Acclimation was studied by measuring leaf osmotic pressure and osmotic pressure at zero turgor. A pattern similar to seasonal leaf water potential emerged. Further, the dryland treatment showed higher osmotic pressure, particularly at zero turgor, during winter and spring. It was concluded that these trees may have acclimated. Diurnal cycles of stomatal resistance, transpiration and leaf water potential on a summer, two winter and a spring day were monitored. Dry land trees showed acclimation, with delayed reaction to environmental water demand and decreasing soil moisture. Trees of the 80 kPa treatment showed greatest stress. Fruit water potential became more negative between April and July, with fruit softening becoming more rapid. Treatment differences were inconclusive. Polyphenol oxidase activity (PPO), soluble and total, was measured. For fruit picked in April and July 1983, the 55 kPa treatment showed the lowest activity and the 80 kPa the highest. Storage at 5,5⁰C for 30 days increased the activity, while fruit softening decreased it. July activity was higher than the April-harvested fruit. The same pattern emerged for fruit harvested in April 1985, although treatment differences were not significant. Rainfall was considerably higher during the fruit development period of 1985 fruit as compared with that of 1983. A significant interaction between restricted container ventilation during ripening and irrigation history was obtained, the 80 kPa fruit showing higher PPO activity than 55 kPa fruit. Ethylene evolution during ripening showed a normal climacteric pattern for 55 kPa and dryland fruit, but a delayed peak for 80 kPa fruit. Fruit calcium concentrations showed rapid changes between 7 and 16 weeks after fruit set thereafter remaining constant to harvest. There were no clear treatment differences. Fruit abscisic acid levels at 50% soft (100% is eating soft) were lowest in 55 kPa fruit, and highest in 80 kPa. A significant correlation between these values and soft fruit PPO activity was found. A preliminary fruit quality prediction model is suggested.
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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