Biological Sciences

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A comparative classification of the sourish-mixed bushveld on the farm Roodeplaat (293 JR) using quadrat and point methods.
(1995) Panagos, Michael David.; Westfall, Ribert Howard.; Zacharias, Peter John Kenneth.; Ellis, Roger.
An area and a point-based technique were used together at each of the same 75 sampling sites (stands), on a Sourish-Mixed Bushveld farm, to collect data for the classification and mapping of the vegetation. Both sets of data were synthesized using the same computer program package and the efficacy of the resulting classifications as well as the efficiency of the two field sampling techniques was compared. Following this, a continuous 7 752 point (1 m apart) transect was carried out, traversing the farm, in order to determine the optimum scales at which to sample Sourish-Mixed Bushveld so as to increase classification efficacy and improve community boundary recognition. The results indicated that (1) the arbitrarily chosen sampling scale of 1:8 000 was too large for "farm-scale" studies; (2) the area-based method proved to be satisfactory in that the classification and vegetation map produced with this method were verified spatially and environmentally; (3) the point-based method was deficient as a classificatory and mapping tool at large scales, since too few species were recorded with this method to make any sense of the classification and mapping of the vegetation was not possible; (4) less time per species was spent using the area-based method but because more species per stand were recorded with this method, the point-based method was quicker per stand; (5) the area-based method was easier to use in dense vegetation and irregular terrain; and (6) the optimum sampling scales for Sourish-Mixed Bushveld, as indicated by the synthesis of the continuous transect data, are about 1:12 000, 1:50 000 and 1:250 000.
Resprouting and multi-stemming and the role of the persistence niche in the structure and dynamics of subtropical coastal dune forest in KwaZulu-Natal province, South Africa.
(2008) Nzunda, Emmanuel F.; Lawes, Michael John.; Griffiths, Megan Elizabeth.
Resprouting is an important means of plant regeneration especially under conditions that do not favour regeneration through seeding such as frequent disturbances, low productivity, unfavourable soil conditions, extreme cold and limited understorey light availability. Sprouts may be advantageous over seedlings because they have higher survival and growth rates than seedlings, since they use resources from parent plants unlike seedlings that have to acquire their own resources. Resprouting is well documented for ecosystems that experience severe disturbances that damage aboveground biomass. For example, resprouting is important for plant persistence against fire in fire-prone savannas and Mediterranean shrub-lands, and hurricanes and cyclones in tropical forests. In these ecosystems, resprouting often results in multi-stemming, because this dilutes the risk of damage among many stems, improving the chances of individual survival. This study was conducted at coastal dune forest at Cape Vidal in north-eastern South Africa, where there is a high incidence of multi-stemmed trees due to resprouting in response to chronic disturbances of low severity. This study examines (1) the importance of resprouting to tree survival and dynamics in an environment where disturbance severity is low but pervasive, and (2) how this resprouting strategy differs from the more familiar sprouting response to severe disturbances such as fire and hurricanes. Analysis of the relationship between multi-stemming and a number of disturbances potentially causing multi-stemming revealed that stem leaning and substrate erosion were the most important disturbances associated with multi-stemming. There were fewer multistemmed trees on dune slacks that had a stable substrate and were protected from sea winds than on dune crests and slopes that had unstable substrate and were exposed to sea winds. Trees resprouted and became multi-stemmed from an early stage to increase their chances of survival against leaning caused by strong sea winds and erosion, and occasional slumping of the unstable dune sand substrate. These low severity disturbances are persistent and are referred to as chronic disturbances in this thesis. As a result of these chronic disturbances, both single and multi-stemmed trees had short stature because taller individuals that emerged above the tree canopy would be exposed to wind damage. Under chronic disturbances plants may manifest a phylogenetically determined sprouting response. However, in this study resprouting and multi-stemming were the results of the tree-disturbance interaction and not a property of a plant or species and were not phylogenetically constrained. Because the disturbances are predominantly of low severity, leaning trees were able to regain the vertical orientation of the growing section by turning upward (a process referred to as ‘turning up’ in this study) and hence survive without resprouting. Species that were prone to turning upward had a low incidence and degree of leaning of their individuals, low frequency of loss of primary stems and high abundance of individuals. Although turning up is less costly to the individual than resprouting, it could only be used by leaning trees that had small angles of inclination and were not eroded. High intensities of the latter require that individuals resprout to survive. The form and function of resprouting varied between seedlings and juvenile and mature trees. Resprouting in seedlings resulted in a single replacement shoot, unlike sprouting in juvenile and mature trees that resulted in multi-stemmed trees. Like sprouting in juvenile and mature trees, sprouting in seedlings was not phylogenetically constrained. Resprouting in seedlings increased seedling persistence; hence species with more sprout seedlings had larger individual seedlings and seedling banks. Resprouting in seedlings increased the chances of seedling recruitment, whereas resprouting in juvenile and mature trees increased the chances of an established plant maintaining its position in the habitat. After disturbances of high severity, which destroy the photosynthesizing parts, plants resprout using carbohydrates stored below- or above ground. In this study, good resprouters stored more carbohydrates both below- and above ground than poor resprouters. The carbohydrates were mobilized for resprouting after disturbance. More carbohydrates were stored in stems than in roots because the prevailing disturbances were mostly of low severity and hence above ground resources were readily available. Similar to storage by plants in severely disturbed habitats, carbohydrates were stored by reserve formation, which competes for carbohydrates with growth and maintenance and forms permanent storage, rather than accumulation, which temporarily stores carbohydrates in excess of demands for growth and maintenance. Stored carbohydrates are not necessary for resprouting of plants after disturbances of low severity because they can resprout using resources remobilized directly from the disturbed photosynthesizing parts. However, in this study, stored carbohydrates served as a bet-hedge against occasional severe disturbances that occurred in addition to chronic disturbances. Allocation of carbohydrates to permanent storage diverts them from growth and reproduction and hence good resprouters had lower growth rates, seed output, seed size and seedling recruitment than poor resprouters. However, the costs of these traits that resulted in low recruitment from seed by good resprouters, were compensated for by high persistence of established individuals of good resprouters through recruitment of sprout stems. This study demonstrates that resprouting is not only advantageous in severely disturbed environments, but also in environments where disturbances are of low severity but nevertheless confer an advantage on individuals that persist. Thus in forest environments where aboveground biomass is seldom destroyed and individuals are relatively long-lived, resprouting can confer significant fitness and selective advantage on individuals.
Intersex in four South African racial groups in Durban.
(1970) Grace, Hatherley James.; Bain, Peter G.
Chapter I The ontogeny of sex development is introduced and a definition of the intersexual state is given. A resume of intersexes recorded in South Africa is presented together with figures showing the increased demand for cytogenetic services in the country. The motivation for this study is discussed. Chapter II The biological concept of sex is outlined together with notes on the evolution of the dioecious state in animals. Thereafter a brief account of the normal and abnormal intersex states in invertebrates and vertebrates is given. Chapter III Examples of some of the abundant references to ambisexual individuals found in the classical literature, history and mythology are listed and the status of intersexes in modern society is discussed: it does not vary much from the times of ancient Greece and Rome. Chapter IV A summary of the events which led to the establishment of cytogenetics as an independent discipline is given. The discovery of nuclear sex dimorphism is noted and methods of investigation of intersexual patients are described. A detailed description is given of cytogenetic technical. "methods, ; and leads to a diagrammatic explanation of the differential diagnosis of intersexes. This is used as a basis for the classification and accordingly a simplified, unambiguous system is proposed and defended. The sources and numbers of pat~ ients recorded in this thesis are explained. Chapter V The sex chromatin is described in detail? with a description of the methods used for its demonstration and evaluation. A pilot study of normal women of the four races established that there is no inter-racial variation in chromatin frequency. Reasons are given for disregarding nuclear appendages as an accurate indication of 'sex '. Chapter VI Embryological stage s i n the development of the human urogenital tracts are recounted after considering the mechanisms of sex determination. Pathogenesis of intersexuality is considered briefly. Chapter VII Gonadal dysgenesis of the female, including Turner's syndrome and phenotypes of both male and female are presented. The classification, pathology, aetiological and cytogenetic factors are described. Illustrative case reports and extensive discussion of the syndrome are presented. Chapter VIII The Klinefelter syndrome occurs in males with supernumerary X chromosomes; a number of chromatin negative, hypogonadal conditions mimic the syndrome: they are described, with notes on the classification, pathology, aetiology and the cytogenetic features. A series of case reports demonstrate both chromatin positive and negative types. Special reference is made to the dermatoglyphic profiles which were discovered to differentiate the XY males into two groups. Chapter IX Three syndromes, the XXX, XYY and agonadism, were not encountered in this study and are consequently not discussed. Chapter X The adrenogenital syndrome has several distinctive signs which are mimicked by induced virilization; these are described and a number of case reports are presented to illustrate the syndromes. Chapter XI Male intersexes occur in several different forms: the classification of the group is explained and a distinction is made between intersexes and those with simple, or 'pseudo-intersexual', defects. The signs of the different syndromes are detailed, together with data concerning the aetiology and cytogenetics. A series of patients' case histories is used to illustrate the various syndromes. Chapter XII Idiopathic female intersexuality is defined and a short explanation of the classification is given. The signs found in this syndrome are detailed and a single case report, which is of exceptional interest because of severe masculinization and the formation of a phallic urethra, is presented. Chapter XIII Hermaphroditism is defined and the subgroups are explained. The variability of signs is discussed and details of anatomy and histology of the genital tract are given . A single hermaphrodite was encountered during the survey and the case is described. Three other patients, all infants, were suspected of being hermaphroditic but because gonadal biopsy was not undertaken they were discharged from hospital without a diagnosis having been made. Discussion of the aetiological mechanisms is presented and the need for early diagnosis and treatment is emphasized. Chapter XIV Some of the simple genital anomalies and somatic defects which resemble intersex are mentioned and are illustrated by attenuated case reports. Chapter XV The incidence of those intersexual conditions recognizable at birth by anatomical deformity is calculated from a total population of 21,000 live births recorded during the calendar year, 1969. The incidence was very low (1:7000) and does not permit analysis of inter-racial distribution. Prevalence is calculated from the number of new patients seen by the author during 1969 in Durban. Some interesting observations were made but the primary objective, to determine whether or not intersexuality is more prevalent in the Bantu than any other race, could not be answered with certainty because of the small number of patients. Hermaphroditism was shown not to be the most common type of intersex syndrome in the Bantu. Chapter XVI An outline is given of the normal dermatoglyphic features of the finger prints and palms. Some references of the patterns in Turner's and Klinefelter's syndromes are available from the literature and are compared with the profiles of patients from this series. A distinct dimorphism was found in chromatin negative hypogonadal males and was thought to distinguish between those with congenital or acquired pathology. Normal values for South Africans have not been established satisfactorily and the urgent need for this information is stressed. Chapter XVII A general discussion of the highlights of this study of the epidemiology, clinical presentation and cytogenetics of intersexuality is given and a number of conclusions are drawn .
Warning vocalisations and predator information transfer in social birds.
(2008) Seoraj-Pillai, Nimmi.; Malan, Gerard.; Slotow, Robert Hugh.
Alarm or warning vocalizations are produced by many animals when they detect a potential predator. However little is known about the information contained in these vocalizations. This study investigated the warning vocalisations of three passerine species, viz. the Buff-streaked Chat Oenanthe bifasciata, the Stonechat Saxicola torquata and the Bronze Mannikin Spermestes cucullatus. The study investigated whether the alarm calls for terrestrial predators differ in their acoustic structure from alarm calls emitted for aerial predators. Birds were exposed to latex terrestrial snakes and mounted aerial raptors, while changes in six acoustic parameters of alarm calls were measured. Bronze Mannikins were investigated for differences in intra-specific alarm calls between familiar and unfamiliar group members by capturing wild groups of mannikins and randomly mixing these to form assorted groups. Bronze Mannikins emitting warning calls were able to discriminate differences in predator size, and increased their calling rate and decreased the end frequency of the alarm call in response to larger predators. This may be the caller’s response to increased threat or variation in frequency may obscure cues to the caller’s whereabouts. Assorted group members were less aggressive to predator models than original members and panicked more during confrontations. Hence the unfamiliarity of the caller may have disrupted group cohesion. The alarm call acoustics of the social Buff-streaked Chats and the solitary Stonechat were compared to examine the effect of group-living on alarm behaviour. Snakes elicited louder calls from both the chat species than raptors. Louder and collectively more vocal social groups might be more successful in discouraging attacks than an asocial species. Variation in amplitude and call frequencies by the Stonechat provided some evidence that they are equally adapted to identifying predator type. Although Buff-streaked Chats increased their call rate in response to nearby predator models, Stonechats produced shorter calls in response to terrestrial predation particularly when the predators were in close proximity. Knowledge gained through direct encounters with predators or the ease with which raptors and snakes can gain access to nests may have played a part in discrimination of predator threat. Overall this study indicated strong correlations between some alarm acoustic parameters and predator size as well as the degree of threat.
Patterns of distribution, diversity and endemism of terrestrial molluscs in South Africa.
(2007) Govender, Vanashrie.; Slotow, Robert Hugh.; Herbert, David Guy.
Molluscs are an important component of South Africa’s biodiversity. The assessment of distribution patterns and factors influencing the biogeographic distribution are an integral part of assessing the conservation status of molluscs and their conservation management needs. The existing terrestrial mollusc data from South Africa were assessed in terms of their value to biodiversity conversation planning and management. Although the data on terrestrial molluscs are incomplete and would be misleading in terms of identifying specific areas for protection, the data do illustrate significant patterns and trends of mollusc endemism and diversity, which can be used to improve biodiversity conservation and management efforts. The distribution of molluscs across the South African landscape illustrated ten broad biogeographical patterns. Two of these patterns reflected ancient distribution patterns of molluscs and consisted of molluscs of the Gondwanaland/southern relict and Laurasian origins. Three biogeographic patterns occurred across the eastern regions. These patterns were defined as the tropical/subtropical east African, subtropical east of southern Africa and east African afromontane patterns. The biogeographic patterns in the west consisted of the characteristic temperate ‘Mediterranean’ Cape centre and the arid regions of northwestern Cape, Namibia and parts of Botswana. An additional biogeographic pattern identified as the nama karoo/central west was recognised. The final two biogeographical patterns described taxa that were widely distributed and taxa that exhibited disjunct distributions. Twenty-six families and forty-three genera were associated with more than one biogeographical pattern. The dominant biogeographic pattern was the tropical/subtropical east African component. Twenty-one families and forty-eight genera were associated with this biogeographical pattern. The east African Afromontane pattern was also a conspicuous biogeographic element in South Africa. Fewer families and genera were distributed in the western and central regions. The distributions of terrestrial molluscs were influenced by a combination of various factors, which included the presence of rivers, the escarpment, altitude, humidity, precipitation, temperature and biomes. Rivers could possibly restrict the distribution of certain mollusc taxa but did not appear to be the dominant factor that influenced the distribution of molluscs across the landscape. In terms of the effect of temperature on the distribution of molluscs, the mean daily and mean annual temperatures appeared to have more of an influence on the distribution patterns than the mean daily maximum and minimum temperatures. Mean annual temperatures influenced the distribution of all families and genera. The mean daily maximum temperature appeared to have little or no effect on the distribution of mollusc taxa. Humidity and biomes also appeared to influence the distribution of taxa. The least inhabited biome was the succulent biome. Many mollusc taxa occurred in the wetter, warmer areas with high humidity levels. Areas of high species richness and high endemic species richness in South Africa were identified using two systems of endemism, one based on distinctive gaps in the frequency distribution of terrestrial molluscs in South Africa and the other based on an existing classification of invertebrate endemism (Hamer & Slotow, 2002). Areas of high mollusc species richness and endemism were also compared to areas of high millipede species richness and endemism. The total number of South African mollusc endemics was 370 (83 % of 447 indigenous species). The dominant mollusc families in South Africa were Achatinidae, Charopidae, Streptaxidae, Subulinidae and Urocyclidae. The first system of endemism identified 56 site endemics (species with only one locality), 50 local endemics (0 < maximum distance < 60 km) and 145 regional endemics (60 km < maximum distance < 330 km). The Hamer & Slotow (2002) classification of endemism classed 67 species as site endemics (maximum distance between localities < 10 km), 47 as local (11 km < maximum distance < 70 km) endemics and 59 as regional endemics (71 km < maximum distance < 150 km). The analysis of mollusc data, with both systems of endemism, showed similar areas of high species and endemic species richness. Quarter-degree grid cells with highest species richness overlapped with grid cells with the highest number of endemic species. However these grid cells coincide with areas that have been intensively sampled and this bias limits the application of the data in conservation planning. The patterns of endemism for molluscs and millipedes within the provinces differed, indicating that the inclusion of a single taxon in conservation planning would inadequately reflect the diversity of invertebrates in South Africa. A preliminary list of specific priority endemic sites for terrestrial mollusc conservation was identified. It is essential that the existing data on invertebrates be evaluated and used to identify key patterns and trends in invertebrate diversity as this will allow for the inclusion of invertebrates in biodiversity conservation planning and management. The analysis of the existing mollusc data identified bio geographical patterns that are important to conservation planning both at the local and national level as well as commonalities and differences between molluscs and millipede distributions. The analysis also highlighted the importance of municipal areas for conservation of hotspots of diversity, particularly in the eastern coastal areas of South Africa.
The physiology of pinus patula seedlings in response to water stress and the implications for plantation regeneration in South Africa.
(2008) Rolando, Carol Ann.; Little, Keith MacMillan.; Pammenter, Norman William.
Pinus patula Schiede ex Schlect. & Cham. is the most widely planted softwood species for both pulpwood and saw timber in the South African forestry industry. High mortality of this species, often in excess of 20%, following planting is currently of major concern and has the potential to limit future deployment for commercial timber. Water stress is often reported to be a cause of mortality during regeneration in commercial forestry plantations yet, prior to 2007, there was no published research on the water relations of P. patula during regeneration in South Africa. This, together with questions raised by the industry as to the role of using water in the planting operation, initiated the series of studies conducted for this thesis. Water planting (application of water into the planting hole at the time of planting) of P. patula seedlings has been used commercially to reduce post-planting water stress and buffer against potentially extreme weather conditions immediately after planting. However, the primary role of the water, as well as its success in increasing survival following planting, has never been critically assessed. Since the use of water in the planting operation is expensive, it was essential that the benefits to using water were quantified, in terms of survival and growth, and justified, in terms of any monetary investment. In addition, there was a lack of local studies investigating the physiological characteristics of P. patula seedlings, particularly their tolerance to low soil water availability. To understand the role of water during the regeneration of P. patula in terms of plantation management and seedling physiology, a variety of research methodologies were used that included: applied field trials, multivariate methods (a retrospective investigation), pot trials and the development of a simple financial model. Four field trials were implemented to test the response in P. patula survival to water applied at planting. Two trials each were situated in the KwaZulu-Natal (KZN) Midlands and Mpumalanga Escarpment. The first trial at each site was planted in spring (October) and the second in summer (February). Watering treatments consisted of different quantities of water used in the planting operation and included 0.5 litres, 2 litres, 4 litres and no water (dry plant). Only at the spring planted trial in the KZN Midlands was survival of the dry planted seedlings significantly lower than that of the seedlings planted with water, at 90 days after planting. This may have been due to low rainfall during the week before and two weeks after planting, or the small size of the seedlings used in the trial. Application of 0.5 litres of water to the planting pit was sufficient to increase survival to a level equivalent to that where 2 or 4 litres of water was used, yet only increased soil moisture in the area immediately surrounding the seedling. This suggested that the role of the water applied during planting was increased root to soil contact. Overall, these four trials indicated that planting with water had the potential to increase survival only when soil water availability was low and rainfall sporadic. There was no effect of water applied at planting on early tree growth. While the results of the four field trials provided an indication of the effect of planting with water on subsequent survival of P. patula seedlings, there was concern that the results of the four trials may not be a true reflection of a dynamic situation. Survival in response to water applied at planting may vary from year to year and across forestry regions due to the unpredictable nature of rainfall and high air temperatures during the planting season, as well as the wide range of forestry sites across which P. patula seedlings are planted. To improve our understanding, a database of 58 trials was compiled where water and dry planting had been carried out. In this way it was possible to investigate whether the results from the four field trials were reflected in a range of previously conducted field trials implemented across time and space. The trials incorporated into the dataset were all planted to P. patula between 1990 and 2005 in the summer rainfall region of southern Africa. Data related to the climate, local weather, physiography and site management at each trial were also included. Summary statistics, linear correlation and multiple regression were used to determine if site-associated variables were related to an increase in survival in the water relative to the dry planted treatments. The analyses indicated that for all 58 trials, survival was lowest during the summer months, regardless of planting treatment. Planting with water was most likely to increase survival when used during spring, autumn and winter planting, although (as with the four applied field trials) there was no overall significant relationship between water planting and survival. Based on these results it was anticipated that an understanding of the water stress physiology of P. patula seedlings was required to explain the observed trends from a more fundamental perspective; if planting with water did not always increase survival, why not? Three pot trials were conducted to increase the understanding of the water relations of P. patula seedlings. These trials were also used to provide benchmark physiological data related to stressed (water) and unstressed seedlings. The first pot trial highlighted the importance of root plug moisture at the time of planting for increasing subsequent survival. The subsequent two pot trials were aimed at investigating the interaction between planting stock quality (as determined by measures of size) and soil water availability and the effect on survival, growth and physiology of P. patula seedlings. These results indicated that P. patula seedlings were not as sensitive to high air and soil temperatures (above 30°C) and low soil water availability (below -1.5 MPa) as previously thought. The seedlings were able to tolerate low soil water availability for several weeks and, following rewatering, were able to recover from moderate and severe water stress (a shoot water potential of below -1.5 MPa). This data supported the results from the four applied field trials and retrospective study of 58 trials, where the application of water to the seedlings at planting did not substantially increase survival. In the pot trials, stomatal conductance started to decrease when shoot water potential approached -0.8 to -0.9 MPa. Stomatal closure occurred at a shoot water potential between -1.2 MPa to -1.5 MPa. Mortality due to water stress occurred only in response to extended periods of low soil water and was associated with a shoot water potential of below -3.0 MPa. There was variability between seedlings in their potential for survival and growth. Inherently bigger seedlings had a greater capacity for new root growth following planting. New root growth, as well as a greater mass of new roots, was associated with higher shoot water potentials and higher rates of transpiration under conditions of low soil water availability. This indicated that seedling quality, as determined by size, may play a role in sensitivity to water stress. The field trials, retrospective study and pot trials indicated that the practice of planting with water was not always critical to the survival of P. patula seedlings. A simple financial model was developed to estimate whether planting with water represented a cost that could be used as a decision criterion, given certain growth parameters and management scenarios. The data projected by the model were also compared to actual research data for water versus dry planting (and the inclusion of an insecticide in the water). While these comparisons were specific to the parameters included in the model for this study, as well as the results of the research trials used in the benchmarking exercises, the model indicated that; 1) costs for planting with water were likely to be recovered only when no blanking (replacing of dead trees) was carried out, with capital invested at a low return rate (3%), 2) including an insecticide in the water increased the likelihood of cost recovery, and 3) site quality had an impact on the increase in survival required to recover planting method costs, with a greater percentage increase in survival required on lower quality sites. Lower quality sites often have a lower mean annual precipitation (associated with higher rainfall variability), or shallow soils (associated with lower soil water availability) and therefore are also likely to be sites where foresters may want to use water to reduce (drought related) mortality. The impact of site quality is thus also an important factor to include in any decisions regarding planting methods (i.e. using water) and their costs. Further investigations should be aimed at examining; 1) the interaction of root plug size (as determined by container type) and soil water availability on growth and physiology of P. patula seedlings, 2) the methods of grading seedlings within a population to select those that have a high potential for survival and growth, and 3) the effects of soil water availability on the physiology, survival and growth of P. patula cuttings, as well as other pine species and hybrids grown in South Africa, such as P. elliottii, P. elliottii x P. caribaea and P. patula x P. tecunumanii. It is likely that the proportion of forestry regions planted to these hybrids will increase in the future.
Sink regulation of photosynthesis in sugarcane
(2007) McCormick, Alistair James.; Watt, Derek Alexander.; Cramer, Michael D.
The C4 plant, sugarcane (Saccharum spp. hybrids), accumulates sucrose to high concentrations and, as a result, has been the focus of extensive research into the biochemistry and physiology of sucrose accumulation. Despite this, the relationship between source leaf photosynthetic rates and sucrose accumulation in the culm has not been well documented. The observations that photosynthetic activity declines during culm maturation in commercial cultivars and that high-sucrose accumulating ancestral genoptypes photosynthesize at rates two-thirds of those of low-sucrose ancestral Saccharum species indicate that source-sink communication may play a pivotal role in determining sucrose yield. The relationship between source and sink tissues in sugarcane was investigated using a supply-demand paradigm, an approach novel in the study of the crop. The demand for photosynthate from the primary culm growth sink was shown to be closely linked to photosynthetic rates, sucrose export and the eventual physiological decline of source leaves. Results from initial field experiments revealed that leaf assimilation rates were negatively correlated with leaf hexose concentrations, but not those of sucrose. Further manipulation of leaf sugar status, through sugar-feeding and cold-girdling techniques, demonstrated the regulatory role of leaf sugar concentrations on photosynthetic activity, thus revealing sucrose, and particularly hexose, as key signal molecules in the modulation of the amount of photosynthate available for export to the sink. Gene expression profiling, by means of array technologies, indicated that changes in leaf sugar status and photosynthetic rates result in concurrent modifications in the expression of several genes involved in fundamental metabolic pathways, including photosynthesis, carbohydrate metabolism, stress response and sugar-signaling. Notable amongst these, was the identification of a potential trehalose 6-phosphate (T6P) sugar-signaling mechanism, thus implicating the trehalose pathway as a central regulatory system in the communication of sink carbon requirements to the source leaf. This study demonstrated that maturation of the culm results in a decreased demand for sucrose, which invokes a sugar-mediated feedback signal to decrease leaf photosynthetic supply processes. However, sugarcane leaves appear to retain the capacity to increase the supply of assimilate to culm tissues under conditions of increased assimilate demand. Uncoupling of the signaling pathways that mediate negative feedback between source and sink tissues may result in improved leaf assimilation rates and, consequently, lead to increased sugarcane sucrose yields.
The effect of elevated atmospheric CO2 on the growth and physiology of Chromolaena odorata.
(2008) Lalla, Reshnee.
Rising atmospheric CO2 (Ca) concentrations have generated concern among scientists, mainly because of CO2’s role as a greenhouse gas and its influence on plant growth and development. Previous research has suggested that future CO2 enriched atmospheres may enhance the success of invasive aliens. Chromolaena odorata is an example of an invasive alien proving to be a serious threat to indigenous vegetation in South Africa, and effective control measures are desperately needed to curb infestations in the future. The current study aimed at assessing the response of C. odorata to elevated Ca and interactive factors, and was divided into two trials. During PART A, C. odorata was grown in competition with 2 grass species: Eragrostis curvula and Themeda triandra (selected for their differential preferences to nutrient availability). All three species were potted in a greenhouse at the University of KwaZulu-Natal (Howard College). There were 16 pots in total, and each pot contained four C. odorata plants, four T. triandra seedlings, and four E. curvula seedlings. Eight pots were exposed to elevated Ca (~700ppm), and eight pots were exposed to ambient Ca (~370ppm). The pots at each Ca treatment were further divided: four received high nutrient treatments (3L per addition), while the other four received low nutrient treatments (300 ml per addition). Studies on growth (e.g. plant height, dry weight, etc.), as well as physiology (e.g. Jmax), were undertaken. Results showed that generally, plants responded positively to high nutrient treatments. In contrast, elevated Ca did not affect growth or any of photosynthetic parameters of C. odorata significantly, but did reduce stomatal limitations. During PART B, C. odorata plants were grown monospecifically to assess whether there was a “chamber effect” associated with planting density. Pots at both Ca treatments contained either four C. odorata or two C. odorata seedlings. Growth and physiology were assessed. The fact that elevated Ca did not affect any of the photosynthetic parameters studied, suggests that photosynthetic down-regulation did not occur. This, together with the fact that no increase in stomatal limitations were observed in elevated Ca, implies that enhancement of photosynthetic assimilation could have occurred in C. odorata plants exposed to CO2 enrichment. Results from this study (PART A and PART B), when compared to previous research on this species, suggests that CO2 enrichment may enhance the success of monoculture populations of C. odorata. However, other species may gain competitive advantages over C. odorata occurring in mixed communites, under CO2 enriched environments. In addition, results of this study support the prediction that increasing Ca will reduce the importance of carbon as an external limiting resource, and that the extent of a plant’s response to Ca enrichment will depend on resources other than CO2. If increases in temperature caused by elevated Ca increases nutrient availability in the soil, then Ca could indirectly enhance the success of C. odorata occurring in mixed communities.
Environmental stochasticity and African elephant population dynamics : investigating limitation through juvenile mortality.
(2008) Woolley, Leigh-Ann.
The successful conservation management of African elephants depends largely on understanding the fundamental processes driving the population regulation of this species. Southern Africa’s increasing populations have raised concern over the impact of high elephant densities on the system, in stark contrast against the elephant’s more precarious position in other parts of Africa. As we search for solutions from the processes of historical elephant regulation, we realise that there is a decided lack of empirical evidence to explicitly direct our efforts. In this PhD, I attempt to investigate the application of the classic pattern of large herbivore population limitation, which mainly involves high juvenile mortality in response to stochastic environmental events, to African elephant population dynamics. Firstly, I evaluated the magnitude and frequency of mortality events that would be required to prevent elephant population growth. The death of 85 % of infants and weaned calves would need to occur twice a generation, while a single severe mortality event (causing the death of all infants and weaned calves and 10 % of the rest of the population) once a generation would be sufficient. However, the severity of these events is not matched in natural occurrence in Africa today and only a single recorded event in Tsavo National Park, Kenya, in the 1970’s has come close when more than 7 000 died during a very severe drought. Secondly, I evaluated the potential role of fire as a stochastic, massmortality event limiting elephant populations. I found that fire functions in a similar manner to other environmental catastrophes and primarily causes high juvenile mortality. However, this catastrophic event also highlighted the extreme behavioural and physiological impacts experienced by the elephant population involved. The potential role of these types of events on long-term female fecundity needs further investigation. In isolation, this type of mortality event would need to occur with high frequency to prevent population growth. However, in combination with a decrease in female fecundity, these stochastic events may have a much greater impact on population demography than first thought. Thirdly, I investigated a potential mechanistic link between stochastic mortality events and juvenile susceptibility to resource limitation. Allometric relationships dictate that juveniles select a diet of higher quality than adult elephants. We found that this was achieved by weaned calf selection of higher quality plant parts, although use of plant types and plant species was similar to that of adult females, who they move across the landscape with. The strong sexual dimorphism exhibited by this species was reflected in adult male use of lower quality forage than adult females (or juveniles) in both dry and wet seasons. Diet quality scaled negatively with body size, but adult females consistently selected a higher quality diet than adult males, irrespective of body size. The nutritional and reproductive demands placed on an individual during different life-history stages therefore influence foraging strategies, together with nutrient requirements, e.g. phosphorus for pregnancy/lactation selected consistently by females when unrestricted in the wet season, protein for growth selected consistently by weaned calves. Competitive displacement of adult females to feed at higher levels in the canopy by calves also influenced feeding behaviour. Therefore intraspecific body size, nutritional requirements (in terms of nutrients and energy) and competition had a strong influence on foraging strategy employed by age-sex classes of elephants in response to seasonal environmental change. More selective juvenile foraging requirements means that juveniles are most susceptible to resource limitation, for example during stochastic environmental events such as droughts. In small, closed systems, juvenile mortality is likely to have a strong influence on elephant population regulation, with a slight, temporary decrease in female fecundity possibly acting in conjunction with juvenile mortality effects. Therefore, stochastic environmental events such as drought and fire may be the only natural incidence of population regulation to occur in these systems, where populations continue to grow exponentially and there is no evidence of density-dependence (as in the case of many small, fenced reserves in South Africa). In large, open, high-density systems in other parts of southern Africa, density dependence acts strongly on female fecundity and causes low levels of juvenile mortality in areas of local population aggregation. Therefore, in isolation, natural juvenile mortality is unlikely to regulate African elephant populations, but in conjunction with decreased female fecundity in response to density-dependent feedbacks and stochastic environmental events, population regulation may occur. The management of long-lived megaherbivore species with similar demographic drivers must include an appreciation of the complexity of population response to manipulation of mortality or fecundity effects. Small changes can potentially result in large shifts in population dynamics. Further insight into the mechanisms driving these processes will allow sound scientific support of megaherbivore management decisions to be made throughout Africa.
The ecological implications of sea-level rise and storms for sandy beaches in KwaZulu-Natal.
(2008) Harris, Linda R.
The aesthetic appeal of beaches has made coastal properties prime sites for development. However, this development has been mismanaged and is within the littoral active zone. Beaches retreat landwards as sea levels rise, but with current development trends, beaches are trapped in a coastal squeeze. Climate-change predictions include an increase in frequency and heightened intensity of storms, which can cause significant erosion. This study aimed to determine the ecological implications of sea-level rise and storms for beaches in KwaZulu-Natal (KZN), using geographic information systems (GIS) and beach sampling methods. The beaches were mapped in terms of physical and biological attributes. Spatial trends in these attributes showed that the coastline can be split into three – the northern, central and southern regions. Although 25 % of the coastline is protected by marine reserves, these are located in the Delagoa bioregion: 28 macrofauna species in the Natal bioregion are not protected. Storm impacts for beaches can be heterogeneous, depending on local coastal features, e.g., nearshore reef and sand dunes, and represented a temporary disturbance to macrofauna communities. A GIS-based coastal recession model was derived from Bruun’s rule, and applied for different scenarios of sea-level rise and coastal development. Coastal squeeze is concern, particularly in the southern region. Further, the 10-m elevation contour was not completely effective as a setback line, even for a low sea-level rise scenario. The coastal recession model was validated using data from a real event in KZN, where sea level rose temporarily by ~1.0 m. The model performed well, although the calibration possibly did not span a wide enough range of beach morphodynamic types, and under-predicted retreat for dissipative beaches. It was concluded that the Natal bioregion needs marine reserves, and that higher resolution spatial data are required for accurate beach modeling and the south coast railway line should be relocated proactively. Guidelines for sandy beach systematic conservation planning were outlined, and seated in a conceptual framework of managing beaches for resilience. Application of the proposed recommendations and frameworks could aid in determining a way forward in integrated coastal zone management for KZN, in the face of the uncertainties associated with climate change.
Behaviour of bottlenose dolphins : inference for dolphin tourism off Durban, South Africa.
(2008) Mahomed, Maleeha.
During long-term studies of dolphins, the number of individuals in the population being studied are constantly monitored using the technique of photo-identification. This constant monitoring makes use of different researchers over time. Therefore, measurement of photographic quality and individual distinctiveness for photo-identification analyses was incorporated in this dissertation to provide an additional data set for analysis. Researchers with differing levels of experiences did not obtain the same information from the same photograph and were unable to reliably quantify variables of photo quality and individual distinctiveness, but experienced researchers were found to be more adept than inexperienced researchers in counting notches on the dorsal fin of bottlenose dolphins. These results highlight the necessity for researchers to be trained in photo-identification techniques prior to carrying out their study. This study theodolite tracked dolphins off Durban from June 2004 to Feb 2005 to assess habitat utilization of bottlenose dolphins (Tursiops aduncus) in the Durban bay region. Dolphins were seen in all months surveyed and throughout the survey area. Temporal distribution of dolphins was skewed with 91% of dolphins seen before midday and 98% of sightings observed in water depths less than 30m. Six behavioural categories were recorded, including: social, fast travel, slow travel, feeding, resting and milling. The most dominant behaviour exhibited by dolphins was slow travel (46%) followed by feeding (27%). Resting was not observed at all. Of the feeding behaviour 88% occurred in the southern end of the bay whereas other behaviours occurred randomly throughout the survey area. Additional theodolite tracks were conducted during experimental boat approaches (before, during and after boat approaches) to determine potential short-term reactions of dolphins to dolphin watching boats. Two speeds of approach (slow ~ < 5 km/hr and fast ~ > 40 km/hr) and two distances of approach (20m and 80 m) were tested. The bottlenose dolphin groups did not change their behaviour in response to boat approaches during any of the periods of experimentation. Short-term changes in group speed, group size and spread were not statistically significant. Dolphin groups continued with their ‘normal’ behaviour and spent the same amount of time in the bay when compared to their distribution and behaviour in the absence of the experimental boat. These findings indicate that the experimental boat did not affect the behaviour of dolphins at either a slow or fast approach and even at a close distance. This is interpreted as being as a result of habituation of the dolphins due to their residency in a busy port. This work is crucial in developing guidelines for the development of a sustainable dolphin watching industry off Durban.
Physiological and cytological biomarker studies using Perna perna for marine pollution monitoring.
(2009) Rajkumar, Anisha.; Smit, Albertus J.
Urbanised and industrial coastal areas in South Africa are most vulnerable to the effects of marine pollution, and the Kwa-Zulu Natal coastline is particularly at risk. The mussels Perna perna, from a polluted (Isipingo Beach) and unpolluted (Park Rynie) site, and Brachidontes semistriatus were evaluated for their use as potential bioindicator organisms. The mussels were subjected to increasing copper concentration treatments to asses the following biomarker responses: cardiac activity, lysosomal membrane stability, malate dehydrogenase enzyme (MDH) activity and body condition index. Brachidontes semistriatus exhibited significant variations in biomarker responses only when exposed to higher Cu dosages, whereas P. perna from Park Rynie displayed distinct changes in heart rate, lysosomal membrane stability and MDH activity with increasing contaminant exposure. Perna perna from Isipingo Beach displayed significant biomarker variation in cardiac activity and lysosomal membrane stability, however differences in MDH activity were only evident at the highest Cu concentration of 100 mg.L-1. Both species from the different Cu treatments failed to show any significant changes in body condition indices due to the limited time of contaminant exposure. The mussel P. perna was therefore selected as a suitable biomonitoring species, and cardiac activity, lysosomal membrane stability and body condition index were chosen as reliable biomarkers for the study. Native P. perna from KZN responded to a distinct pollution gradient along the coastline by displaying significant bradycardia, reduced lysosomal membrane stability, poor condition indices and high heavy metal tissue concentrations. Durban, Isipingo and Umkomaas were singled out as the most contaminated sites along the coast, and Zinkwazi and Park Rynie as the least polluted. In addition, significant correlations between tissue and sediment metal concentrations suggest that the species is an effective heavy metal bioaccumulator of Cd, Pb, Zn, Cr and Fe. Similar spatial trends in seawater and mussel tissue concentrations in Durban Harbour were identified. Stations in the port located nearest to the freshwater inputs and stormwater drains displayed the highest metal concentrations in tissue and seawater, as well as adverse biomarker responses from transplanted P. perna. These results suggest that Durban Harbour is strongly influenced by tidal exchange and contaminated freshwater inflow entering the harbour. It was also found that reproduction imposes a considerable effect on P. perna body condition as spawning events in winter months result in pronounced body mass loss. The study concluded that P. perna is a highly effective bioindicator species, and cardiac activity, lysosomal membrane stability and body condition index can successfully be employed in marine pollution monitoring programmes.
The performance and rooting of eucalyptus grandis x nitens cuttings.
(2007) Murugan, Nelisha.; Watt, Maria Paula Mousaco Deoliveira.; Mycock, David John.; Mokotedi, Mompe Edward Oscar.
Hybrid clones of Eucalyptus grandis and E. nitens (GN) have consistently been shown to be suitable for planting in cold, dry, marginal plantation sites, where they exhibit high yields and superior pulp properties. However, their clonal propagation is hindered by the very poor rooting success of cuttings. The present study aimed at assessing the effect of cutting type, time of year of setting cuttings and Seradix 2 application on rooting and development of cuttings of a commercially important Eucalyptus grandis x Eucalyptus nitens clone (GN107). Cuttings were prepared from clonal hedge coppice at the Mondi Business Paper, Trahar Technology Centre, Hilton. Three cutting types were used (cut at different distances from the node) for each terminal (situated below the apical bud) and non-terminal cuttings. The leaves were trimmed and, for half the cuttings, the base of the stem of cuttings were dipped in Seradix 2 rooting powder (3 g kg-1 4-(indole-3-yl)-butyric acid (IBA). They were then placed into rooting trays (128 inserts/ tray arranged as 8 rows x 16 columns). Seradix 2-treated and Seradix 2-untreated terminal and non-terminal cuttings, cut at, above and below the node (twelve treatments in total) were set in trays with one treatment per column of eight replicates, per tray. There were nineteen trays overall. The trays were filled with peat, perlite and vermiculite (3:3:1) and were maintained in a Mondi greenhouse, with air temperature at 25°C to 27°C (thermostatically activated fans), root zone temperature at 28°C (bed heaters) and 20 second misting at 10 minute intervals (automatic misters). The study was carried out in November 2005, April 2006 and June 2006. In the first experiment, both terminal and non-terminal cuttings were used; thereafter only non-terminal cuttings were used. The plantlet yield was very low, regardless of cutting type, Seradix 2 treatment and the time of year the cuttings were set. The highest plantlet production (12.5%) and rooting frequencies (13.8%) were achieved with non-terminal cuttings treated with Seradix 2. Although not statistically significant, Seradix 2 inhibited shoot production (31.4% for Seradix 2-untreated and 24.2% for treated cuttings). The position at which inserts were cut in relation to the node did not significantly affect the number of plantlets produced and non-terminal cuttings appeared hardier and performed better than terminal cuttings. The time of year of setting cuttings did not have any significant effect on plantlet yield, nonetheless, plantlet yield was highest in cuttings set in November (9.2%) and lowest in April (0.4%). In addition, cuttings set in November (spring), had superior shoot development in terms of the number of cuttings that produced shoots (regardless of root production), shoot length and the mass of shoots relative to root mass. The highest percentages of cuttings that produced roots (regardless of shoot growth) (10%) and the highest number of roots per cutting (2) were part of the June trial. Therefore, cuttings set in June (winter) had superior root development as compared with cuttings set in November (spring) or April (autumn). In all of the studies, three rooting patterns were observed in cuttings: roots produced only from the cut area only (type 1), only from the sides of the stem (type 2) and from both sites (type 3). Non-terminal cuttings treated with Seradix 2 showed a higher incidence of types 2 and 3 rooting patterns than the terminal cuttings. Seradix 2 application increased the prevalence of types 2 and 3 rooting patterns. Although not statistically different, cuttings dipped 2.5 cm into Seradix 2 produced more types 2 and 3 rooting patterns than cuttings dipped at the abaxial end only. Light microscopy of stem sections of cuttings indicated that roots appeared to originate from the xylem archs as well as from the cambium. The collected data indicate that it is necessary to continue research towards improving the efficiency of plantlet production of GN107 via cuttings. It appears that cuttings of this clone may be set throughout the year and that terminal cuttings should be avoided. In addition, the present practice at the Mondi Hilton nursery of treating cuttings with Seradix 2 needs to be reconsidered as although it increases rooting, it does not increase plantlet production due to its apparent inhibitory effect on shoot development.
Seed germination and medicinal properties of Alepidea species.
(2009) Mulaudzi, Rofhiwa Bridgeht.; Van Staden, Johannes.; Finnie, Jeffrey Franklin.
The rhizomes of Alepidea amatymbica and Alepidea natalensis are used for medicinal purposes. Because of the increase in demand for these plants the species is becoming scarce. As the seed biology of neither species is well defined, conditions as well as treatments required for optimum germination and vigour were studied. Seeds were exposed to various physical factors such as varying light and temperature conditions and cold stratification, sowing depth and seed storage. The effects of smoke-water, butenolide (3-methyl-2H-furo [2, 3-c] pyran-2-one) a novel smoke compound and chemical substances (gibberellins, kinetin and KNO3) were also tested in order to improve seed germination. Alepidea amatymbica and A. natalensis achieved the highest seed germination (72.5% and 80%, respectively) at 25 °C under a 16 h photoperiod with a mean germination time (MGT) of 18 and 12 days, respectively. Phytochrome studies showed that A. natalensis requires light for germination. Cold stratification (5 °C) for 14-28 days significantly improved the percentage germination of both species (> 90%) compared to non-stratified seeds (control) at 25 °C under a 16 h photoperiod. Sowing A. amatymbica and A. natalensis seeds at a depth of 0.5 cm resulted in higher percentage germination compared to 2.5 cm. The highest emergence rate for A. amatymbica was 40% at a sowing depth of 0.5 cm and the lowest emergence rate was 3% at 2.5 cm. Six months storage of A. natalensis seeds at room temperature (25 ± 2 °C) showed maximum germination (99%) with a MGT of 9 days. Smoke-water treatment of A. amatymbica seeds significantly enhanced germination from 72% to 91%. Smoke and butenolide at 10 °C and 25 °C promoted germination of A. natalensis seeds in a 16 h photoperiod. Smokewater application significantly improved both germination and seedling vigour of A. natalensis. GA3 (10-8 M) was the best treatment for achieving maximum percentage germination of A. natalensis seeds. Antibacterial (two Gram-positive bacteria: Bacillus subtilis, Staphylococcus aureus and two Gram-negative bacteria: Escherichia coli, Klebsiella pneumoniae), antifungal (Candida albicans), anti-inflammatory (COX-1 and -2) and genotoxicity tests (Ames test) were carried out on petroleum ether (PE), dichloromethane (DCM), 80% ethanol (EtOH) and water extracts of the two Alepidea species. Water extracts of A. natalensis rhizomes exhibited high activity (MIC values of 0.78 mg/ml) against the four bacterial strains. High activity was also observed in the PE and DCM leaf extracts of the same plant against the Gram-positive bacteria. The PE and DCM extracts of A. amatymbica rhizomes exhibited the best activity (MIC values of 0.39 mg/ml) against Bacillus subtilis. The rest of the extracts showed low activity (MIC values >1 mg/ml). All the extracts showed activity against Candida albicans, with A. natalensis leaf extracts exhibiting the highest antifungal activity with MIC values of 0.88, 0.20 and 0.78 mg/ml for PE, DCM and EtOH, respectively. EtOH extracts had inhibition less than 40% for both A. natalensis and A. amatymbica. All the PE extracts showed higher inhibitory activity for COX-2 than for COX-1. PE and DCM extracts had percentage inhibitions above 70% in both COX-1 and COX-2 assays. The Ames test for genotoxicity revealed that none of the plant extracts were genotoxic to the Salmonella TA98 tester strain.
Pharmacology and phytochemistry of South African plants used as anthelmintics.
(2009) Aremu, Adeyemi Oladapo.
Traditional medicine in South Africa is part of the culture of the people and has been in existence for a long-time. Although animal components form part of the ingredients used, plant material constitutes the major component. South Africa is endowed with vast resources of medicinal and aromatic plants which have been employed for treatment against various diseases for decades. A large number of South Africans still depend on traditional medicine for their healthcare needs due to its affordability, accessibility and cultural importance. Helminth infections are among the variety of diseases treated by traditional healers. These infections are regarded as neglected tropical diseases (NTDs) due to their high prevalence among the economically disadvantaged living in rural areas in different regions of the world.
Pharmacological, phytochemical and safety evaluation of commercial herbal preparations common in South Africa.
(2009) Ndhlala, Ashwell Rungano.
Herbal formulations claimed to cure several medical conditions including skin eruptions, chest pains, wounds, gout, menstrual pains, stress, nervous disorders, microbial and viral infections as well as stomach ailments have recently appeared as part of South African traditional medicine. The formulations consist of mixtures of extracts of different plant parts from several different plant species packaged in labelled bottles or boxes. The mixtures are available for sale in herbal shops and public places. While there has been widespread use of these herbal mixtures, there has been no scientific evidence to support their use. This project was aimed at documenting, validating claimed efficacy and testing the safety of fourteen unregistered commercial herbal preparations commonly sold in Pietermaritzburg. A detailed investigation of the pharmacological effects and safety of the plant components of one of the mixtures, Imbiza ephuzwato was also carried out.
Micropropagation and medicinal properties of Barleria greenii and Huernia hystrix.
(2009) Amoo, Stephen Oluwaseun.; Van Staden, Johannes.; Finnie, Jeffrey Franklin.
The crisis of newly emerging diseases and the resistance of many pathogens to currently used drugs, coupled with the adverse side-effects of many of these drugs have necessitated the continuous search for new drugs that are potent and efficacious with minimal or no adverse side-effects. The plant kingdom is known to contain many novel biologically active compounds, many of which could potentially have a higher medicinal value when compared to some of the current medications. Indeed, the use of plants in traditional medicine, especially in African communities, is gaining more importance due to their affordability and accessibility as well as their effectiveness. Exponential population growth rates in many developing countries has resulted in heavy exploitation of our plant resources for their medicinal values. In addition, plant habitat destruction arising from human developmental activities has contributed to the fragmentation or loss of many plant populations. Owing to these factors, many plant species with horticultural and/or medicinal potential have become either extinct or are threatened with extinction. These threatened species cut across different taxonomic categories including shrubs, trees and succulents. Without the application of effective conservation strategies, the medicinal and/or horticultural potential of such threatened species may be totally lost with time. The extinction of such species could lead to the loss of potential therapeutic compounds and/or genes capable of being exploited in the biosynthesis of new potent pharmaceutical compounds.
Systematic studies in Gnidia L. (Thymelaeaceae)
(2010) Beaumont, Angela Jane.; Edwards, Trevor John.; Van Staden, Johannes.
Gnidia L., variously estimated to contain 100–160 species, is the largest genus in the sub-cosmopolitan family Thymelaeaceae. Most species are shrubby, and occur in tropical and southern Africa, with one species reaching southern India and Sri Lanka, and 14 species endemic to Madagascar. Assorted segregate genera have been established using characters considered by some as too few, too trivial or unreliable. Generic limits have been contentious with authors following either a narrower concept of Gnidia or a broader circumscription within which segregate genera are placed in synonymy under Gnidia. Regional treatments for African and Madagascan floras have been published over the last century until very recently, but the genus was last revised in its entirety 153 years ago. Today, a broad-based concept of Gnidia is generally recognised, but there is no modern infrageneric classification, and species relationships are poorly understood. Homogenous groups of species are identified by their similarities of leaf length and width or bract length and width ratios. Species comprising the homogenous groups for leaf ratios differ to those comprising the homogenous groups based on bract ratios, and there is no correlation between leaf and bract length and width ratios. This suggests that the factors influencing leaf diversity differ from those influencing bract diversity. Bracts differ most from leaves in species with capitate inflorescences, and involucres of several layers of bracts likely protect reproductive organs (flowers) in heads. Previously overlooked morphological and micromorphological details, and morphometric analyses of leaf, bract and floral dimension data help define individual species, and clades of species derived from phylogenetic analyses of molecular data. Evidence from a phylogenetic analysis of nuclear ribosomal and plastid DNA sequence data confirms the polyphyly of Gnidia. Three lineages contain Gnidia species and species of genera from southern Africa, southern South America or Australia, while another lineage corresponds largely to the previously recognized genus Lasiosiphon. The genus Lasiosiphon is reinstated characterised by flowers mostly in heads, bracts different from the leaves, and the presence of smooth hairs; it now includes species with tetramerous flowers as well as ones with pentamerous flowers. Gynodioecy is recorded for the first time in a single species and represents the first documented example of sexual polymorphism involving unisexual flowers in Gnidia and sub- Saharan Thymelaeaceae. The findings of this thesis are discussed in terms of their phylogenetic value and contribution to our better understanding of the generic limits of Gnidia and its relationships with other southern hemisphere Thymelaeoideae. The circumscription and generic affinities of Gnidia as suggested by results presented in this thesis are compared to previous classification systems for congruence and dissimilarity.
Regulation of hyperhydricity in Aloe polyphylla propagated in vitro.
(2009) Ivanova, Mariyana Vasileva.; Finnie, Jeffrey Franklin.; Van Staden, Johannes.
Micropropagation of Aloe polyphylla, an endangered species with a high ornamental and medicinal value, is an important part of its conservation. However, the in vitro culture was hindered by the phenomenon of hyperhydricity. The research reported in this thesis was undertaken for two reasons. Firstly, to understand the role of various culture factors involved in the process of hyperhydricity in A. polyphylla and to identify the in vitro conditions, under which this disorder can be prevented. Secondly, we conducted an investigation into the underlying mechanisms of this phenomenon by probing if it was mediated through internal cytokinins. Ammonium (NH4 +) ions, applied cytokinins (CKs) and CK concentrations were tested in multifactorial combinations and significantly influenced the regeneration rate and occurrence of hyperhydricity. Shoots were grown on media with different NH4 + concentrations (10.3, 20.6 and 61.8 mM) and supplemented with BA, zeatin or TDZ at 0, 5 or 15 ìM. Elevating the levels of NH4 +, in the absence of CKs, could not induce hyperhydricity. Similarly, very low hyperhydricity was observed when CKs were added to media containing low NH4 + (10.3 mM). However, in the presence of higher NH4 + concentrations, CKs increased hyperhydricity in a concentrationdependant manner, suggesting that they were capable of inducing this syndrome only when other factors in the culture system were not optimised. High numbers of healthy looking shoots were produced on media with low NH4 + and low BA or zeatin (5 ìM). The use of TDZ resulted in the formation of buds, which did not develop into shoots. In view of the fact that NH4 + was supplied in the form of NH4NO3, it was difficult to determine if NH4 + or nitrate (NO3 -) ions were associated with the increase in hyperhydricity. We further examined the role of nitrogen (N) supplied as inorganic NH4 + or NO3 -, or organic glutamine. The omission of total N from the culture medium resulted in low multiplication and hindered shoot growth. Ammonium as the sole source of N depressed shoot regeneration and growth and escalated the frequency of hyperhydricity to ca. 50%. When NO3 - was used as the sole N source, shoots of fine quality were produced and hyperhydricity was completely eliminated. Overall, the MS N mix was superior to any single N source for multiplication and growth of shoots, suggesting a synergistic effect between NH4 + and NO3 - on shoot regeneration. Furthermore, not only the absolute amount of N, but also the relative amounts of NH4 + and NO3 - influenced the multiplication rate, frequency of hyperhydricity and shoot quality. The highest regeneration was obtained with NH4 + : NO3 - ratios (mM) of 20 : 40, 30 : 30 and 40 : 20. Decreasing the ratio of NH4 + : NO3 - lowered the occurrence of hyperhydricity. The potential of glutamine as the sole source of N was also demonstrated, since its application resulted in the production of good quality shoots and almost no hyperhydricity. Shoot explants grown in static liquid media became hyperhydric and lost their ability to regenerate. The type of gelling agent used to solidify the medium affected greatly hyperhydricity and shoot multiplication. Gelrite resulted in a significantly lower multiplication rate and four times higher hyperhydricity (64.7%) compared to when agar was used. Gelrite was further selected to test the hypothesis if hyperhydricity can be overcome by decreasing the relative matric potential of the media, and respectively the availability of water, as represented by increasing gelrite concentrations. Satisfactory reduction in hyperhydricity was achieved only at 16 g l-1 gelrite, however the regeneration also decreased. The nature of the gelling agent is therefore essential for the successful control of this phenomenon. It appears that a crucial prerequisite for the reduction of hyperhydricity in tissue cultures of A. polyphylla is the gaseous exchange between the in vitro atmosphere and the outside environment. In ventilated cultures, achieved by using a modified lid with a hole (d = 7 mm) covered with polyester or cotton mesh, hyperhydricity was completely eliminated, irrespectively of the type of gelling agent. Ventilation was further advantageous for the in vitro regenerants by increasing their leaf chlorophyll content as well as epicuticular wax deposition, the last one being indicative of the development of the water loss regulation mechanisms of explants. The increased culture ventilation, however, was negatively correlated with the regeneration rate and shoot growth. Endogenous CKs were measured in in vitro regenerants after an eight-week cycle to examine whether the hyperhydricity-inducing effect of exogenous CKs and gelling agents is associated with changes in the endogenous CK content. The content of endogenous CKs, determined by HPLC-mass spectrometry, in the shoots grown on CK-free media comprised isopentenyladenine-, trans-zeatin- and cis-zeatin-type CKs. The application of exogenous CKs resulted in an increase in the CK content of the shoots. Following application of zeatin, dihydrozeatin-type CKs were also detected in the newly-formed shoots. Application of BA to the media led to a transition from isoprenoid CKs to aromatic CKs in the shoots. Shoots grown on gelrite media contained higher levels of endogenous CKs compared to those on agar media. Total CK content of hyperhydric shoots was higher than that of normal shoots grown on the same medium. We suggest that the ability of exogenous CKs and gelrite to induce hyperhydricity in shoots of Aloe polyphylla is at least partially due to up-regulation of endogenous CK levels. However, hyperhydricity is a multifactor process in which different factors intervene.
Heavy metals in South African medicinal plants.
(2008) Street, Renée Anne.; Van Staden, Johannes.; Stirk, Wendy Ann.; Southway, Colin.
Plants are able to take up and accumulate certain environmental contaminants such as heavy metals. When the plants are ingested by man, these contaminants are transferred along the food chain. Due to the poorly regulated medicinal plant trade in South Africa, many opportunities exist for heavy metal contamination of medicinal plants namely contaminated harvest sites as well as poor drying, processing, storage, transport and manufacturing conditions. The concentrations of five heavy metals (As, Cd, Co, Ni, Pb) and six microelements (B, Cu, Fe, Mn, Mo, Zn) were determined in some commonly used South African medicinal plants obtained from street markets. Elemental content was determined using inductively coupled plasma optical emission spectrophotometry (ICP-OES). Some of the medicinal plant samples investigated contained As and Cd at levels exceeding the World Health Organization limits of 1 and 0.3 mg kg-1 respectively. Lead and Ni were detected in all the samples. Elevated Fe and Mn levels were recorded in certain plant species. The results revealed multiple metal contamination in some medicinal plant parts sold in local markets and is thus grounds for concern. The effects of Cd application on growth parameters of some medicinal plant species belonging to the Hyacinthaceae (Albuca setosa, Eucomis autumnalis, Eucomis humilis, Merwilla plumbea) gave insight into heavy metal accumulation and distribution in these species. Application of Cd at 5 mg l-1 over a 12 week period reduced growth in A. setosa. The medicinally used A. setosa bulbs accumulated 37 mg kg-1 Cd after 12 weeks. Cadmium application at 2 mg l-1 over a six week period had no effect on growth parameters of E. autumnalis or E. humilis. However, a substantial difference in total Cd accumulation was detected in the plants (40.2 and 15.3 mg kg-1 respectively). Cadmium application at 2 mg l-1 significantly reduced the fresh weight of leaves, bulbs and roots of M. plumbea. Although most of the Cd was stored in the roots, the medicinally used bulbs accumulated up to 11.6 mg kg-1 when applied at 10 mg l-1. The antagonistic effect between Cd and Zn treatments and their effect on micronutrient distribution in M. plumbea were investigated. Five treatments were evaluated: (1) Hoagland’s nutrient solution (HS) (control) (2) HS + Cd 2 mg l-1 (single) (3) HS + Cd 2 mg l-1 + Zn 50 mg l-1 (combination) (4) HS + Cd 2 mg l-1 + Zn 100 mg l-1 (combination) (5) HS + Cd 2 mg l-1 + Zn 150 mg l-1 (combination). Cadmium readily accumulated in leaves, bulbs and roots of M. plumbea when supplied at 2 mg l-1. Zinc at 50 mg l-1 led to increased Cd accumulation. However, further increases in Zn concentration showed an antagonistic effect of Zn on Cd uptake and accumulation. Thus, increasing Zn levels in soils may be favourable for reducing toxic Cd accumulation in M. plumbea plants. Boron was not significantly affected by the addition of Cd to the media. However, with an increase in Zn, leaf B content increased while the B content in the bulbs and roots decreased. Copper and Mo levels were not significantly affected by treatments with Cd or Cd/Zn combinations. Compared to the control, Cd and Cd/Zn applications caused an increase in Mn content in leaves, bulbs and roots. Iron levels of M. plumbea were not significantly affected by Cd in the media. However, with an increase of Zn in the Cd-containing media, Fe content in the leaves, bulbs and roots increased. Tulbaghia violacea is one of the few medicinal plants that is also frequently used as a leafy vegetable. Application of Cd at 2 and 5 mg l-1 to T. violacea of varying sizes (small 8 - 10 g, medium 16 - 20 g, large 80 – 95 g) elicited a difference in growth response, Cd accumulation and micronutrient distribution. Leaf length and fresh weight of leaves of the medium-size plants decreased with application of Cd at 2 mg l-1 whilst 5 mg l-1 Cd significantly decreased the number of leaves in small-sized plants. Small plants accumulated more Cd in the leaves than medium- or large-sized plants. Application of Cd at 2 mg l-1 and 5 mg l-1 lowered the leaf Cu, Fe, Mo and Zn contents in small- and medium-size plants. This study indicated that T. violacea has the ability to accumulate Cd. In addition, plant size plays an important role with regards to Cd accumulation and elemental distribution. The effect of various nutrient applications (10%, 50% and 100% Hoagland’s nutrient solutions (HS); and HS deficient in N, P or K) on growth parameters and micronutrient distribution in Dioscorea dregeana were investigated. Irrigating plants with 50% HS resulted in better growth performance, whereas a deficiency of either N, P or K negatively affected seedling growth. Plants grown in 10% HS contained higher total B, Fe and Mo levels compared to seedlings grown in 50% and 100% HS. Compared to the control, P deficiency resulted in a Fe increase in the leaves, tuber and roots while a lack of P and K significantly increased total Mn content in D. dregeana. The effect of excess Zn (100, 200 and 300 mg l-1) on growth performance, chlorophyll content and microelemental distribution on Dioscorea sylvatica was investigated. Growth parameters showed a significant decrease when supplied with Zn at 100 mg l-1. Zinc phytotoxicity was evident by the reduction in chlorophyll content. Highest Zn concentrations were detected in the roots. Certain micronutrients appear to be redistributed due to Zn toxicity. The effect of microelements (Cu, Zn) and heavy metals (Cd, Pb, Hg) on germination and seedling development of Bowiea volubilis, Eucomis autumnalis and Merwilla plumbea was investigated. Copper and Zn applied at 1 mg l.1 significantly reduced the percentage germination of E. autumnalis. Low concentrations (. 1 mg l.1) of Cu and Zn negatively affected the root growth of all three species. Mercury concentrations of 0.5 and 1 mg l.1 significantly decreased the percentage germination of B. volubilis and E. autumnalis respectively. Cadmium and Hg at 2 mg l.1 showed a negative effect on the root growth of B. volubilis. Concentrations of 0.5 mg l.1 of all heavy metals tested significantly decreased shoot length of M. plumbea. The effect of Cd on biological activity (anti-inflammatory, antibacterial and antifungal) of medicinal plants with previously confirmed activity was evaluated. When supplied with Cd at 2 mg l-1, Eucomis humilis bulbous extracts showed lower anti-inflammatory activity than the control for both COX-1 and COX-2 activity. Eucomis autumnalis bulbous extracts had greater COX-1 activity compared to the control. However, Cd suppressed the activity of COX-2. Compared with non-Cd-treated Merwilla plumbea plants (control), those supplied with Cd at 10 mg l-1 showed increased antibacterial activity against Bacillus subtilis, Klebsiella pneumoniae and Staphylococcus aureus. However, no change in activity against Escherichia coli was observed. Cadmium accumulation in the bulbs had no effect on antifungal activity of Tulbaghia violacea. Thus, optimized agricultural practices are essential for quality control of cultivated medicinal plants. The studies presented in this thesis collectively answer several questions related to heavy metal involvement in South African medicinal plants. The findings substantiate the need to regulate and monitor the South African medicinal plant trade against heavy metal contamination which will in turn provide a product of safety and quality to the consumer.

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