Doctoral Degrees (Botany)

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Cape elements on high-altitude corridors and edaphic islands.
(2004) Carbutt, Clinton.; Beckett, Richard Peter.; Edwards, Trevor John.
Common to the temperate floras throughout sub-Saharan Africa is a group of taxa with strong ties to the Cape Floristic Region (CFR) (≈ Cape elements). Their distribution is limited to the eastern escarpment of Africa (e.g. the Drakensberg Alpine Centre - DAC), on nutrient-rich humic soils, as well as on isolated sandstone outcrops of low elevation, on nutrient-poor soils (e.g. the Pondoland Centre - PC), suggesting that intrinsic soil fertility is not the primary determinant of their distribution. The principal aim of this study was to determine which aspect of the edaphic environment of the DAC is most influenced by temperature, that may indirectly render it nutrient-poor and therefore provide suitable niches for Cape elements, as in the PC. A multidisciplinary approach involving aspects of plant biogeography, plant ecology, plant ecophysiology and soil chemistry was therefore adopted. The study regions were the DAC, PC and the KwaZulu-Natal Midlands. The flora of the DAC was resurveyed for this study, and is richer than previously thought: 2818 native taxa, most of which (2520) are angiosperms. The phytogeography of the DAC and PC is discussed, and comparisons are made with the floras of KwaZulu-Natal and the CFR. Their climatic environments, as well as those for the CFR and Sneeuberge, were compared using rainfall and temperature data from a range of sources. These climatic regimes were correlated with the floristic patterns of Cape elements for the high-altitude regions of South Africa and Lesotho. Altitude and rainfall increased, and temperature decreased, as the number of Cape elements increased towards the DAC. This study provided a contemporary inventory of the Cape elements of the DAC and PC. A total of 89 genera are recognised as Cape elements, of which 60 (c. 67%) are shared between the two regions. The highest number of Cape elements recorded for the eastern escarpment was the DAC (72 genera), with the highest number from all sites analysed being the PC (77 genera). The most Cape elements are contributed by the Asteraceae, Scrophulariaceae, Iridaceae, Fabaceae, Orchidaceae and Restionaceae, partly due to the success of annual aerial parts and their geophytic growth forms, which are convergent in these families. Further compartmentalisation into life and growth forms shows that most Cape elements of the DAC and PC are either ericoid (and sclerophyllous) or mesic herbs and shrubs. The ecological and ecophysiological aspects of this study involved the use of reciprocal pot experiments established along a gradient of altitude from coastal hinterland to mountain, that investigated the interactions between altitude, temperature and substrate on plant productivity in sites known either to support or to exclude Cape elements. Three soils were used at each site, representative of the DAC, PC and KwaZulu-Natal Midlands. The interactions between 'soil' and 'site' (≈ the climatic environment) were quantified using a temperate test taxon (Diascia) that has a strong Cape-centred distribution. Plant characters relating to morphology and nutrient content, and soil characters relating to fertility, were used as the basis for comparing treatment effects (soil-site interactions). Soil nitrogen availability was assayed using pot experiments with Eragrostis curvula (Schrad.) Nees. Wheat pot experiments revealed no Al³⁺ toxicity in 'Drakensberg' soil. Non-metric multidimensional scaling (NMDS) and redundancy analysis (RDA) indicated that all soil-site interactions were significant contributors to biomass differences, and that the Cape taxon performed poorly in the nutrient-rich Drakensberg soil at low altitude. Soil samples indicated that Drakensberg soil was the most nutrient-rich, and Pondoland soil the most nutrient-poor. Although total nitrogen in Drakensberg soil was six times higher than Pondoland soil, both soils mineralised similar low levels of nitrogen at their respective spring temperatures. The result for Drakensberg soil (simulated so as to include the effect of altitude) meant that only 1.7% of its total nitrogen was mineralisable at 12°C (its mean spring temperature). These findings suggest that nitrogen mineralisation rate is a key growth-limiting factor in the DAC, exacerbated by a number of complex interactions with soil pH and organic matter. It is hypothesized that Cape elements are preadapted to high-altitude habitats. These habitats are nutrient-deprived due to low temperatures, which reduce metabolic rates and the movement of ions in cold soils. This constraint imposes nutrient-related stresses similar to those of the CFR and PC. Taxa that are adapted to the nutrient-poor soils of the CFR are preadapted to the temperature-induced 'nutrient-poor' soils of the DAC and vice versa. This 'compatibility' has allowed the reciprocal exchange of taxa between regions, as suggested by cladistic biogeographical analyses using Cliffortia, Disa, Moraea and Pterygodium. The strong overlap of Cape elements between the CFR and PC is a product of similar nutritional niches and ancient floristic continuity. The result therefore is a high number of Cape elements common to the DAC and PC.
The influence of vegetation control on the growth and pulping properties of a eucalyptus grandis x camaldulensis hybrid clone.
(1999) Little, Keith MacMillan.; Van Staden, Johannes.; Clarke, G. Peter Y.; Roberts, Peter J. T.; Beckett, Richard Peter.
In order to determine if weed control as practised during the establishment phase of tree growth had a beneficial and long term (over a six to eight year rotation) impact on tree performance, a Eucalyptus hybrid clone (GC304) was planted in a field trial in 1990. The trial was situated in the coastal Zululand region near the KwaZulu-Natal town of Mtunzini. Nine different vegetation management treatments were imposed from establishment. These included a weedy control, a manually weeded treatment, a chemically weeded treatment, a 1.2 m row and 1.2 m inter-row weeding, a 0.5 m radius ring weeding, a complete weeding except for a 0.5 m radius ring around the tree, and the use of two legume cover-crops, Mucuna puriens (cowpea) and Vigna sinensis (velvet bean). Initial improvements in tree performance from these competition control treatments were detected from 60 days after planting, and were maintained over seven growing seasons. This occurred despite the absence of competitive vegetation after the first growing season due to reduced light, following crown canopy closure. There were strong indications that initial competition was mainly for moisture and possibly also for nutrients, rather than competition for light. Initially those trees that had weeds within their immediate vicinity were most affected (weedy control, inter-row weeding and the complete weeding except for a 0.5 m radius around the tree). With time, tree performance was more closely related to an increase in the percentage of the area kept free of weeds. The best performing treatment at felling, the manually weeded control, produced 17.1 % and 38.5 % more merchantable timber than the 1.2 m row weeding and the weedy control, at an increased profit of 8 % and 27 %, respectively. Two forms of competition (interspecific and intraspecific competition) were evident in the weedy control at different stages of tree development in contrast to the one (intraspecific competition) in the manually weeded treatment. Interspecific competition resulted in greater variability between the trees in the weedy control by the time canopy closure had occurred. This differentiation in tree size was further enhanced by asymmetric intraspecific competition once the trees had become established. The onset of intraspecific competition was first detected 995 days after planting for the manually weeded treatment and 1641 days after planting for the weedy control. Of the various competition indices that were tested in order to try and explain this differential growth in terms of individual tree performance, none was able to do so to complete satisfaction. The growth rates of different tree size classes were therefore compared for the weedy control and manually weeded treatment. The diverging slopes of the different stem area classes indicated that the larger trees were growing at the expense of the smaller trees. This type of competition is known as asymmetric intraspecific competition. In addition, a comparison was made between the slopes for the weedy and weedfree treatments for similar stem area classes. No significant difference was detected, indicating that similar size classes in these two treatments grew at similar rates. Trees from three treatments were selected (manually weeded treatment, 1.2 m row weeding treatment and the weedy control) and tested for the wood and pulping properties of density, active alkali consumption, extractable content, screened pulp yield, pulp yield per hectare and fibre length and coarseness. The use of Canonical Variate Analysis to determine if there were differences between the three treatments for the variates measured, indicated that they were significantly different. There was a significantly positive trend of an increase in density, extractable content and active alkali consumption with increased weed control. A possible explanation for this could be that the larger trees of the manually weeded treatment were under more stress (from increased intraspecific competition) during the latter phase of their growth. This was demonstrated by comparing the growth rates for these three treatments. The smaller trees of the 1.2 m row weeding treatment and the weedy control exhibited a lower rate of decline. As no significant difference was detected for screened pulp yield between the treatments, any differences in the pulp yield per hectare values could be attributed to differences in the merchantable volume. There was a 22.6 % and 40.8 % increase in the pulp yield per hectare for the manually weeded treatment in comparison to the 1.2 m row weeding treatment and the weedy control. The planting of cover-crops, although beneficial in terms of weed suppression, caused significant tree suppression. This occurred despite the fact that their initial biomass accumulation was slower than that of the natural weed population. Of the two cover-crops, the use of the velvet bean was not considered suitable due to its vigorous vining habit which affected the growth form of the trees. Subsequent work suggests that if the beneficial qualities of cowpeas are to be realised (that of weed suppression, erosion control and nitrogen fixation), a delay in their planting by three months after establishment of the trees should alleviate any negative impacts on tree growth.
Mechanisms of desiccation tolerance in cryptogams.
(2002) Mayaba, Nosisa.; Beckett, Richard Peter.
In this study adaptations of bryophytes and lichens to desiccation stress were examined. The aim was to test whether desiccation tolerance in the selected species is constitutive or if desiccation tolerance could be induced by various hardening treatments. In addition, some putative tolerance mechanisms were investigated, including the accumulation of sugars, increase in ROS scavenging systems and other mechanisms e.g. energy dissipating processes. To determine if hardening treatments prior to desiccation stress increased desiccation tolerance, mosses and lichens were partially desiccated or treated with ABA. The effect of hardening treatments on the physiology of the moss Atrichum androgynum and lichens Peltigera polydactyla, Ramalina celastri and Telochistes capensis during a desiccation-rehydration cycle was investigated. Photosynthesis, respiration and chlorophyll fluorescence measurements were used as rapid tools to determine the metabolic activities in these lichens and moss species. In A. androgynum partial desiccation following slow drying at 52% RH increased the rate of recovery of net photosynthesis. Net photosynthesis recovered almost completely following slow drying in the material that was partially dehydrated and/or treated with ABA. This suggests that partial dehydration hardens the moss, and that ABA can fully substitute for partial dehydration. In R celastri and P. polydactyla both partial dehydration and ABA treatments displayed some improvement in desiccation tolerance depending on the duration and severity of stress. The reduction in the re-saturation respiration burst in P. polydactyla, although not quite significant, strongly suggests that hardening increases mycobiont tolerance. However, it is more difficult to establish whether the hardening treatments improve photobiont performance. In the moss A. androgynum ABA treatment increased the rate of recovery of photosynthesis and PSII activity, and also doubled non-photochemical quenching (NPQ). Increased NPQ activity will reduce ROS formation, and may explain in part how ABA hardens the moss to desiccation. In ABA treated, but not untreated mosses, desiccation significantly increased the concentration of soluble sugars in A. androgynum. Sugar accumulation may promote vitrification of the cytoplasm and protect membranes during desiccation. Starch concentrations in freshly collected A. androgynum and R. celastri were only c. 40 and 80 mg g ¯¹ dry mass respectively, and slightly rose during desiccation, but were only slightly affected by ABA pretreatment. ABA did not reduce chlorophyll breakdown during desiccation. In P. polydactyla ABA pretreatment had little effect on any of these parameters. Changes in the activities of the free radical scavenging enzymes ascorbate peroxidase, catalase and superoxide dismutase were measured during wetting and drying cycles in the moss A. androgynum and in the lichens P. polydactyla, R. celastri and T capensis. These species normally grow in the understorey of the Afromontane forest, moist, xeric, and extremely xeric miicrohabitats respectively. In A. androgyum, enzyme activity was measured shortly after collection, after 3 d storage following hardening by partial dehydration and/or 1 h treatment with ABA or distilled water and during desiccation and rehydration. In A. androgynum enzyme activities of CAT and SOD in untreated material were always higher than in the hardened treatments, while both partial dehydration and ABA treatments tended to reduce both CAT and the induction of SOD activity, although these effects were not significant between the treatments. This suggests that ABA may not be involved in the induction of free radical scavenging enzymes and probably these enzymes are not important in desiccation tolerance of A. androgynum. In lichens, the enzyme activity was measured shortly after collection, after hydration for 48 hat 100% RH, after desiccation for 14 d and 28 d, and during the first 30 min of hydration with liquid water. Enzyme activities tended to rise or stay the same following rehydration in all the species tested. After desiccation for 14 d, enzyme activities decreased, and then decreased further to very low values after 4 weeks desiccation. In all species, including T capensis from an extremely xeric habitat, the activities of all enzymes remained at very low values during the 30 min following rehydration, and were therefore unavailable to remove any reactive oxygen species accumulating in lichen tissues as a result of desiccation. Results suggests that the enzymic antioxidants are more likely to be involved in removing reactive oxygen species produced during the normal metabolic processes of lichens than having a role in desiccation tolerance. The Afromontane understorey moss Atrichum androgynum displayed an oxidative burst of H₂O₂ during rehydration following desiccation. Maximum rates of H₂O₂ production occur during the first 15 min of rehydration. While the production of H₂O₂ increases with increasing desiccation times, the moss produced significant amounts of H₂O₂ during rehydration after desiccation for times that did not inhibit photosynthesis or cause K⁺ leakage. A. androgynum may produce more H₂O₂ during desiccation than rehydration, because desiccation artificially induced using polyethylene glycol strongly stimulates production. Experiments involving inhibitors and exogenously supplied reductants indicate that peroxidases are responsible for the synthesis of H₂O₂. Factors that influence the rate of H₂O₂ production during rehydration include light and the hormone ABA. Patterns of H₂O₂ production are discussed in terms of their possible role as a defence against pathogenic fungi and bacteria.
Occurrence and properties of the multicopper oxidases laccase and tyrosinase in lichens.
(2012) Laufer, Zsanett.; Beckett, Richard Peter.
The work presented in this thesis describes the occurrence and properties of two multicopper oxidases derived from lichens. Despite numerous data on laccases and tyrosinases in fungi and flowering plants, this is the first report of the occurrence of these enzymes in lichenized ascomycetes. Extracellular laccase and tyrosinase activity was measured in 50 species of lichens from different taxonomic groupings and contrasting habitats. Out of 27 species tested from suborder Peltigerineae, all displayed laccase and tyrosinase activity that correlated to each other, while activity was absent in species tested from other lichen groups. Identification of the enzymes as laccases and tyrosinases was confirmed by the ability of lichen thalli or leachates to readily metabolize substrates such as 2,2’-azino(bis-3-ethylbenzthiazoline-6-sulfonate) (ABTS), syringaldazine and o-tolidine in case of laccase and L-dihydroxyphenylalanine (L-DOPA), Ltyrosine and epinephrine in case of tyrosinase in the absence of hydrogen peroxide. The activities of both enzymes were highly sensitive to cyanide and azide, and tyrosinase activity was sensitive to hexylresorcinol. Laccase activity had typical pH and temperature optima and an absorption spectrum with a peak at 614 nm. Tyrosinases could be activated by sodium dodecyl sulphate (SDS) and had typical tyrosinase molecular masses of approx. 60 kDa. The diversity of laccase isoforms in 20 lichen species from suborder Peltigerineae was investigated. The molecular masses of the active forms of most laccases varied between 135 and 190 kDa, although some lichens within the family Peltigeraceae had laccases with higher masses, typically varying from 200 to over 350 kDa. Most species contained one oligomeric laccase isoform. Desiccation and wounding stimulated laccase activity, while only wounding stimulated tyrosinase activity. The ability of laccases to decolorize dye is a classic attribute of laccases, and one with biotechnological potential. The ability of eight lichen species to decolourize different types of dyes was therefore tested. Interestingly, results showed that not only species belonging to suborder Peltigerineae but also species from other lichen group effectively decolourised dyes after 48 h suggesting that other oxidases appear to have ability to decolorize. Hopefully, our work could contribute to the better knowledge and application of lichen multicopper oxidases.
Responses to chilling and cryopreservation of recalcitrant seeds of Ekebergia capensis from different provenances.
(2020) Bharuth, Vishal Behary Lal.; Beckett, Richard Peter.; Pammenter, Norman William.
Recalcitrant seeds are shed at relatively high water contents and are metabolically active. The effect of chilling the recalcitrant seeds of purportedly a single species, Ekebergia capensis, from Port Elizabeth (PE; Eastern Cape), St Lucia (KwaZulu-Natal [KZN]) and Tanzania (tropical southern Africa) was tested. Viability and axis ultrastructure, solute leakage and protein synthesis were investigated. Additionally, cryopreservation of embryonic axes (explants) was studied. In particular, the ability of cathodic water to improve explant survival was tested, and related to its effect on the production of potential harmful reactive oxygen species such as superoxide (·O2-) and hydrogen peroxide (H2O2), and its ability to maintain levels of total aqueous antioxidants (TAA). Shoot and root ultrastructure were examined after each cryopreparative step in explants treated with and without cathodic water, and ultrastructure correlated with survival. The seeds from PE retained 80% viability after 12 weeks storage at 1° and 3°C and 100% when stored at 6°C. Those from St Lucia were dead after 38 d storage at 3°C. All the seeds from Tanzania were dead after 9 d when stored at 3°C. The rate of protein synthesis decreased gradually over the storage period, irrespective of the provenance. Electrolyte leakage from axes showed that those from St Lucia and Tanzania ‘leaked’ solutes irrespective of the duration or temperature or storage, but those from PE showed an initial increase, which then decreased. A 30% mass loss was achieved after 48 d, 230 h and 21 h for seeds from PE, St Lucia and Tanzania, respectively. Nuclear ribosomal ITS1 sequences revealed the presence of three well-to-strongly-supported monophyletic clades corresponding to the geographical areas from which the seeds were sampled (PE, KZN and Tanzania). Axes from the seeds from St Lucia and Tanzania lost ultrastructural integrity during storage while those from PE did not. The levels of ·O2- and H2O2 increased gradually after each cryopreparative step. Using cathodic water allowed 30% of PE explants to survive cryopreservation, while none of the St Lucia explants did. Root ultrastructure was well preserved, however, gradual ultrastructural deterioration was observed in the shoot meristem.

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