Studies of the biology and ecology of the high shore South African limpet, Helcion pectunculus (Mollusca : patellogastropoda)
- Authors: Gray, David Richard
- Date: 1997
- Subjects: Mollusks -- South Africa Limpets -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5776 , http://hdl.handle.net/10962/d1005464
- Description: Investigations were carried out into aspects of the bio!ogy and ecology of Helcion pectunculus along the coait of South Africa. These included studies of the distribution, density and biomass of the limpet at six sites along the east coast plus one site on the west coast; the growth of H pectunculus on both west and east coasts; a comparison of the reproductive biology of the east and west coast populations; the foraging activity and feeding behaviour of H pectunculus and the driving forces behind the rhythmic behaviour of this limpet; the importance of the crevice environment in the biology and ecology of H pectunculus. Helcion pectunculus has a restricted zonation, with the majority of animals residing in crevices in the upper Balanoid zone during diurnallowtides, although individuals were occasionally found in the lower Balanoid zone on shores with a gently sloping aspect. This limpet occurs in higher densities (50-lO0 individuals/m2) on shores which have large numbers of crevices and boulders i.e. quartzitic sandstone shores. On most shores, the ratio of males to females differed significantly from a 1: 1 ratio with the highest ratio being obtained on the west .. coast (3 males: 1 female). At all sites, the populations of H pectunculus exhibited strong sexual dimorphism. Males and females were always found to differ in size, with individuals of < 20 mm shell length generally being male whilst limpets with a shell length of> 22 mm were generally female. Helcion pectunculus grows allometrically, increasing in height faster than length, which is expected of a high shore gastropod mollusc attempting to reduce evaporative water loss. Growth rates were similar on both east and west coasts regardless of the differing oceanographic conditions. The theoretical values of Lmax were also similar being 30.86 mm and 30.71 mm respectively. Micro-growth bands are laid down within its' shell which have the same periodicity as the tidal cycle and these enabled age estimates to be made. Younger individuals were male whilst older animals were female, suggesting that H pectunculus is a protandrous hermaphrodite. Histological examination proved, unequivocally, that this limpet undergoes a protandric sex change, changing from male to female when they are about 2 years old. Both east and west coast popUlations had a marked reproductive cycle, exhibiting two spawning periods a year, one in April and another in November. The possibility that the reproductive pattern exhibited is now phylogeneticallyconstrained is discussed. It is suggested that H pectunculus has evolved a reproductive cycle which will allow its planktonic larvae to utilise the valuable phytoplankton bloom food source whilst using onshore winds to ensure that larvae are not transported out to sea and lost. The number of foraging excursions carried out by individuals of H pectunculus was found to have a significant effect on Gonad Index and hence potential reproductive output. The activity pattern of H pectunculus varied depending upon micro-habitat; animals inhabiting both east and west facing rock surfaces are active during nocturnal low tides whilst animals on west facing rock surfaces are also active during daytime low tides whilst in the shade. Limpets travel further during foraging excursions in winter (X = 85.53 cm) than in either spring (x = 55.7 cm) or summer (X = 48.8 cm) and also during spring low tides (x = 89.8 cm) compared with neaps (x = 40.9 cm). This limpet exhibits rigid homing to a fixed scar within a-crevice and feeding excursions were found to cons.is.t. of three distinct phases, a rapid outward phase, a slower foraging phase and a rapid homeward phase. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rockface with the highest microalgal biomass and also the smoothest rock surface. Helcion pectunculus exhibits a free-running endogenous rhythm of locomotor activity with both circadian and circatidal components and it is suggested that this rhythm plays a role in allowing the limpet to avoid unfavourable environmental conditions. The exogenous entrainment factor of this endogenous rhythm was the time of exposure to air in the field. There was found to be an organized distribution of limpets within crevices with smaller, younger limpets being towards the back of the crevice and larger, older limpets towards the crevice mouth. It is hypothesized that juvenile limpets of this species actively select and settle at the backs of crevices responding to chemical cues of adult conspecifics. The crevice refuge supplies the limpets with a stable and buffered environment with higher relative humidities (X = 72.3%) and lower rock surface temperatures (X = 19.7°C) than adjacent exposed rock surfaces (X = 64.5%; x = 22.9°C). Limpet body temperatures were significantly lower in crevic~refuges compared to limpets on exposed rock surfaces. Body temperatures never exceeded the rock surface temperatures. It is suggested that this is the result of morphological adaptations such as shell ornamentation and allometric growth. Light levels above 1000 J.1E.m-2.s-1 inhibited foraging activity in H. pectunculuswhilst limpets subjected to 30-50% shade foraged even during daytime lowtides. This limpet is one of the least tenacious (2.75 ± 0.13 kg.cm-2 ) of all South African limpets and the possibility that wave activity governs both the activity patterns and homing behaviour of this limpet is discussed. Limpets deprived of a crevice refuge experienced extremely high mortalities, with 45% of the limpets being lost during the first high tide period. A hypothetical model of the hierarchy of exogenous factors controlling limpet foraging activity is introduced and discussed in relation to the results of this study. Finally, it is suggested that in addition to the "migratory" and "non-migratory" groups of limpets present on southern African shores a third group of limpets seem to be present which may be classed as "specialized non-migratory" species. These are species that do not migrate, garden or aggressively fight off like conspecifics. They have overcome the competition for space and food on intertidal rocky shores by adapting to a particular habitat which is exclusive to them alone. From the combined results of these studies, it can be stated that H. pectunculus has adapted physiologically, morphologically and behaviourally to successfully survive the extreme conditions in the upper Balanoid zone.
- Full Text:
- Date Issued: 1997
- Authors: Gray, David Richard
- Date: 1997
- Subjects: Mollusks -- South Africa Limpets -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5776 , http://hdl.handle.net/10962/d1005464
- Description: Investigations were carried out into aspects of the bio!ogy and ecology of Helcion pectunculus along the coait of South Africa. These included studies of the distribution, density and biomass of the limpet at six sites along the east coast plus one site on the west coast; the growth of H pectunculus on both west and east coasts; a comparison of the reproductive biology of the east and west coast populations; the foraging activity and feeding behaviour of H pectunculus and the driving forces behind the rhythmic behaviour of this limpet; the importance of the crevice environment in the biology and ecology of H pectunculus. Helcion pectunculus has a restricted zonation, with the majority of animals residing in crevices in the upper Balanoid zone during diurnallowtides, although individuals were occasionally found in the lower Balanoid zone on shores with a gently sloping aspect. This limpet occurs in higher densities (50-lO0 individuals/m2) on shores which have large numbers of crevices and boulders i.e. quartzitic sandstone shores. On most shores, the ratio of males to females differed significantly from a 1: 1 ratio with the highest ratio being obtained on the west .. coast (3 males: 1 female). At all sites, the populations of H pectunculus exhibited strong sexual dimorphism. Males and females were always found to differ in size, with individuals of < 20 mm shell length generally being male whilst limpets with a shell length of> 22 mm were generally female. Helcion pectunculus grows allometrically, increasing in height faster than length, which is expected of a high shore gastropod mollusc attempting to reduce evaporative water loss. Growth rates were similar on both east and west coasts regardless of the differing oceanographic conditions. The theoretical values of Lmax were also similar being 30.86 mm and 30.71 mm respectively. Micro-growth bands are laid down within its' shell which have the same periodicity as the tidal cycle and these enabled age estimates to be made. Younger individuals were male whilst older animals were female, suggesting that H pectunculus is a protandrous hermaphrodite. Histological examination proved, unequivocally, that this limpet undergoes a protandric sex change, changing from male to female when they are about 2 years old. Both east and west coast popUlations had a marked reproductive cycle, exhibiting two spawning periods a year, one in April and another in November. The possibility that the reproductive pattern exhibited is now phylogeneticallyconstrained is discussed. It is suggested that H pectunculus has evolved a reproductive cycle which will allow its planktonic larvae to utilise the valuable phytoplankton bloom food source whilst using onshore winds to ensure that larvae are not transported out to sea and lost. The number of foraging excursions carried out by individuals of H pectunculus was found to have a significant effect on Gonad Index and hence potential reproductive output. The activity pattern of H pectunculus varied depending upon micro-habitat; animals inhabiting both east and west facing rock surfaces are active during nocturnal low tides whilst animals on west facing rock surfaces are also active during daytime low tides whilst in the shade. Limpets travel further during foraging excursions in winter (X = 85.53 cm) than in either spring (x = 55.7 cm) or summer (X = 48.8 cm) and also during spring low tides (x = 89.8 cm) compared with neaps (x = 40.9 cm). This limpet exhibits rigid homing to a fixed scar within a-crevice and feeding excursions were found to cons.is.t. of three distinct phases, a rapid outward phase, a slower foraging phase and a rapid homeward phase. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rockface with the highest microalgal biomass and also the smoothest rock surface. Helcion pectunculus exhibits a free-running endogenous rhythm of locomotor activity with both circadian and circatidal components and it is suggested that this rhythm plays a role in allowing the limpet to avoid unfavourable environmental conditions. The exogenous entrainment factor of this endogenous rhythm was the time of exposure to air in the field. There was found to be an organized distribution of limpets within crevices with smaller, younger limpets being towards the back of the crevice and larger, older limpets towards the crevice mouth. It is hypothesized that juvenile limpets of this species actively select and settle at the backs of crevices responding to chemical cues of adult conspecifics. The crevice refuge supplies the limpets with a stable and buffered environment with higher relative humidities (X = 72.3%) and lower rock surface temperatures (X = 19.7°C) than adjacent exposed rock surfaces (X = 64.5%; x = 22.9°C). Limpet body temperatures were significantly lower in crevic~refuges compared to limpets on exposed rock surfaces. Body temperatures never exceeded the rock surface temperatures. It is suggested that this is the result of morphological adaptations such as shell ornamentation and allometric growth. Light levels above 1000 J.1E.m-2.s-1 inhibited foraging activity in H. pectunculuswhilst limpets subjected to 30-50% shade foraged even during daytime lowtides. This limpet is one of the least tenacious (2.75 ± 0.13 kg.cm-2 ) of all South African limpets and the possibility that wave activity governs both the activity patterns and homing behaviour of this limpet is discussed. Limpets deprived of a crevice refuge experienced extremely high mortalities, with 45% of the limpets being lost during the first high tide period. A hypothetical model of the hierarchy of exogenous factors controlling limpet foraging activity is introduced and discussed in relation to the results of this study. Finally, it is suggested that in addition to the "migratory" and "non-migratory" groups of limpets present on southern African shores a third group of limpets seem to be present which may be classed as "specialized non-migratory" species. These are species that do not migrate, garden or aggressively fight off like conspecifics. They have overcome the competition for space and food on intertidal rocky shores by adapting to a particular habitat which is exclusive to them alone. From the combined results of these studies, it can be stated that H. pectunculus has adapted physiologically, morphologically and behaviourally to successfully survive the extreme conditions in the upper Balanoid zone.
- Full Text:
- Date Issued: 1997
Synthetic and physical organic studies of chromone derivatives
- Ramaite, Ipfani David Isaiah
- Authors: Ramaite, Ipfani David Isaiah
- Date: 1997
- Subjects: Benzopyrans Heterocyclic compounds -- Derivatives Coumarins
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4380 , http://hdl.handle.net/10962/d1005045
- Description: A range of chromone-2-carboxylic acids has been prepared by condensing suitably substituted 2-hydroxyacetophenones with diethyl oxalate. pK₂ Studies of these acids revealed that 6- or 7-methoxy substituents decreased acidity while the 6-nitro group enhanced acidity; the strongest acid was the 3-chloro derivative, the increase in acidity being attributed to steric inhibition of acid-weakening delocalisation between the carboxyl group and the chromone system. Various chromone-2-carboxamides, derived from acid chloride precursors, were converted to polysubstituted acrylamides by nucleophilic ring-opening with selected amine nucleophiles. The main fragmentation patterns exhibited by these acrylamides were elucidated using a combination of low resolution, high resolution and meta-stable peak analysis, while the effect of substituents on the simultaneous internal rotation involving the carboxamide and enamine moieties were studied using dynamic NMR spectroscopy. Rotational barriers of ca. 67.1 kJmol ̄¹ and ca. 102 kJmol ̄¹ were found for the enamine and amide rotors, respectively. Several synthetic pathways were followed to prepare a series of 2-(N,N-dialkylamino)chromones which were subjected to detailed mass spectral analysis. In addition to substituent-specific fragmentations , the 2-aminochromones appear to fragment via 3 major pathways. The effect of substituents on the internal rotation of the amino moeity was investigated by variable temperature ¹H NMR spectroscopy and the resulting DNMR data was used to calculate the rotational barriers. Examination of the data reveals that the electron-releasing 6- and 7- substituents reduce the C-NMe₂ rotational barrier to ca. 43.5 kJmol ̄¹ , while the nitro analogue has the largest rotational barrier (ca. 46.1 kJmol ̄¹) because of the electron-withdrawing effect of this substituent.
- Full Text:
- Date Issued: 1997
- Authors: Ramaite, Ipfani David Isaiah
- Date: 1997
- Subjects: Benzopyrans Heterocyclic compounds -- Derivatives Coumarins
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4380 , http://hdl.handle.net/10962/d1005045
- Description: A range of chromone-2-carboxylic acids has been prepared by condensing suitably substituted 2-hydroxyacetophenones with diethyl oxalate. pK₂ Studies of these acids revealed that 6- or 7-methoxy substituents decreased acidity while the 6-nitro group enhanced acidity; the strongest acid was the 3-chloro derivative, the increase in acidity being attributed to steric inhibition of acid-weakening delocalisation between the carboxyl group and the chromone system. Various chromone-2-carboxamides, derived from acid chloride precursors, were converted to polysubstituted acrylamides by nucleophilic ring-opening with selected amine nucleophiles. The main fragmentation patterns exhibited by these acrylamides were elucidated using a combination of low resolution, high resolution and meta-stable peak analysis, while the effect of substituents on the simultaneous internal rotation involving the carboxamide and enamine moieties were studied using dynamic NMR spectroscopy. Rotational barriers of ca. 67.1 kJmol ̄¹ and ca. 102 kJmol ̄¹ were found for the enamine and amide rotors, respectively. Several synthetic pathways were followed to prepare a series of 2-(N,N-dialkylamino)chromones which were subjected to detailed mass spectral analysis. In addition to substituent-specific fragmentations , the 2-aminochromones appear to fragment via 3 major pathways. The effect of substituents on the internal rotation of the amino moeity was investigated by variable temperature ¹H NMR spectroscopy and the resulting DNMR data was used to calculate the rotational barriers. Examination of the data reveals that the electron-releasing 6- and 7- substituents reduce the C-NMe₂ rotational barrier to ca. 43.5 kJmol ̄¹ , while the nitro analogue has the largest rotational barrier (ca. 46.1 kJmol ̄¹) because of the electron-withdrawing effect of this substituent.
- Full Text:
- Date Issued: 1997
Termination of psychotherapy : a phenomenological exploration of the therapist's and the patient's experience
- Authors: Schlodder, Michelle Ingrid
- Date: 1997
- Subjects: Psychotherapy -- termination Psychotherapy
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:3049 , http://hdl.handle.net/10962/d1002558
- Full Text:
- Date Issued: 1997
- Authors: Schlodder, Michelle Ingrid
- Date: 1997
- Subjects: Psychotherapy -- termination Psychotherapy
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:3049 , http://hdl.handle.net/10962/d1002558
- Full Text:
- Date Issued: 1997
The effect of elevated CO₂ on Phaseolus vulgaris L. cv Contender
- Authors: Mjwara, Jabulani Michael
- Date: 1997
- Subjects: Plants -- Effect of carbon dioxide on Kidney bean
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4215 , http://hdl.handle.net/10962/d1003784
- Description: The response of Phaseolus vulgaris L. cv. Contender grown in controlled environmental conditions, at either ambient or elevated (360 and 700 μmol mol ̄¹, respectively) CO₂ concentrations ([CO₂]), was monitored from 10 days after germination (DAG) until the onset of senescence. Elevated CO₂ had a pronounced effect on total plant height (TPH), leaf area (LA), dry weight (DW) accumulation and specific leaf area (SLA). All of these were significantly increased by elevated [CO₂] with the exception of SLA, which was significantly reduced. Except for higher initial relative growth rates (RGR) in CO₂-enriched plants, RGR did not differ significantly between the two CO₂ treatments throughout the remainder of growth period. While growth parameters clearly differed between CO₂ treatments, the effects of CO₂ on many physiological processes including net assimilation rate (NAR), Rubisco activity, and some foliar nutrient concentrations were largely transient. For example, CO₂ enrichment significantly increased NAR, but from 20 DAG onward, NAR declined to levels measured on plants grown under ambient CO₂. Similarly, the decline in both foliar N concentration and Rubisco activity in CO₂-enriched plants after 20 DAG was significantly greater than the decline observed for ambient CO₂ plants. Soluble leaf protein and total chlorophylls (a+b) were also significantly reduced in plants grown under elevated CO₂. Chlorophyll (a/b) ratios increased with time underelevated CO₂, indicating that the rate of decline of chlorophyll b was higher than that of chorophyll α. No significant changes in total carotenoid (x+c) levels were observed in either CO₂ treatment. Under enhanced CO₂, the foliar concentrations of K and Mn were increased significantly, while P, Ca, Fe and Zn were reduced significantly. However, changes in Mg and Cu concentrations were not significant. High CO₂-grown plants also exhibited pronounced leaf discoloration or chlorosis, coupled with a significant reduction in leaf longevity. The levels of non-structural carbohydrates (sucrose, glucose, fructose and starch) and nitrogenous compounds (nitrogen, total soluble proteins and free amino acids) were determined for leaves and developing seeds of P. vulgaris. Leaf tissue of elevated CO₂-grown plants accumulated significantly higher levels of both soluble sugars and starch. Leaf ultrastructure revealed considerable erilargement of starch grain sizes with surface areas more than five times larger compared to those of control plants. No apparent differences in structure and membrane integrity of chloroplasts in both CO₂ treatments were noted. Although ambient CO₂-grown plants had comparatively low levels of non-structural carbohydrates (NSC), they accumulated significantly higher levels of nitrogenous compounds. The levels of NSC were consistently higher in seeds of plants grown under elevated CO₂. In comparison to plants grown at elevated [CO₂], pods and seeds of ambient CO₂-grown plants had significantly larger pools of free amino compounds and N. Stomatal conductance (gs) declined significantly, as expected for plants grown under elevated CO₂. This was accompanied by a decline in transpiration rates (E). Reduced gs and E led to high AlE ratio, which meant improved water use efficiency (WUE) values for CO₂-enriched bean plants. Leaf carbon isotope discrimination (∆) against the heavier isotope of carbon (¹³C), has been used to select for high WUE in C₃ plants. In plants grown at elevated CO₂ concentration, ,1 was significantly reduced. Although ∆ was negatively correlated with WUE in both CO₂ treatments, the correlation was steeper and highly negative for CO₂-enriched plants. These results indicate underlying differences in gas-exchange physiology, including stomatal responses between ambient and elevated CO₂-grown plants. Photosynthetic acclimation was investigated using the response of assimilation to internal carbon dioxide concentration (A/C₁ curves). At early stages of growth, the initial slope of the A/C₁ response curve did not differ with CO₂ treatment. In contrast, CO₂-saturated photosynthetic rate (Amax) was significantly higher in plants grown under elevated versus ambient CO₂ at 15 DAG. However, at subsequent stages of growth both the initial slope and Amax declined in bean plants grown in elevated CO₂. Apparent carboxylation efficiency (ACE, estimated from the initial slope of A/C₁ response) values followed a similar trend and were significantly reduced in CO₂-enriched plants. These results indicate that acclimation or negative adjustment of photosynthesis may have been caused by a combination of both stomatal and biochemical limitations. Bean plants grown under conditions of elevated atmospheric CO₂ flowered 3 to 4 days earlier, and produced significantly more flowers and pods than plants grown at ambient conditions. Plants grown at elevated CO₂ aborted 22 and 20% more flowers and pods, respectively, than plants grown at ambient CO₂. Elevated CO₂ also significantly increased the number of tillers or lateral branches produced by plants, which contributed to a significant increase in pod number and seed yield in these plants. Although plants grown at elevated CO₂ produced on average 8 seeds per pod, while plants grown under ambient CO2 conditions produced 5 seeds per pod, the greater number of seeds was offset by lower seed weights in plants grown under _ elevated CO₂. Thus, despite high seed yield in beans grown under elevated CO₂, the harvest index (HI) did not change significantly between CO₂ treatments.
- Full Text:
- Date Issued: 1997
- Authors: Mjwara, Jabulani Michael
- Date: 1997
- Subjects: Plants -- Effect of carbon dioxide on Kidney bean
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4215 , http://hdl.handle.net/10962/d1003784
- Description: The response of Phaseolus vulgaris L. cv. Contender grown in controlled environmental conditions, at either ambient or elevated (360 and 700 μmol mol ̄¹, respectively) CO₂ concentrations ([CO₂]), was monitored from 10 days after germination (DAG) until the onset of senescence. Elevated CO₂ had a pronounced effect on total plant height (TPH), leaf area (LA), dry weight (DW) accumulation and specific leaf area (SLA). All of these were significantly increased by elevated [CO₂] with the exception of SLA, which was significantly reduced. Except for higher initial relative growth rates (RGR) in CO₂-enriched plants, RGR did not differ significantly between the two CO₂ treatments throughout the remainder of growth period. While growth parameters clearly differed between CO₂ treatments, the effects of CO₂ on many physiological processes including net assimilation rate (NAR), Rubisco activity, and some foliar nutrient concentrations were largely transient. For example, CO₂ enrichment significantly increased NAR, but from 20 DAG onward, NAR declined to levels measured on plants grown under ambient CO₂. Similarly, the decline in both foliar N concentration and Rubisco activity in CO₂-enriched plants after 20 DAG was significantly greater than the decline observed for ambient CO₂ plants. Soluble leaf protein and total chlorophylls (a+b) were also significantly reduced in plants grown under elevated CO₂. Chlorophyll (a/b) ratios increased with time underelevated CO₂, indicating that the rate of decline of chlorophyll b was higher than that of chorophyll α. No significant changes in total carotenoid (x+c) levels were observed in either CO₂ treatment. Under enhanced CO₂, the foliar concentrations of K and Mn were increased significantly, while P, Ca, Fe and Zn were reduced significantly. However, changes in Mg and Cu concentrations were not significant. High CO₂-grown plants also exhibited pronounced leaf discoloration or chlorosis, coupled with a significant reduction in leaf longevity. The levels of non-structural carbohydrates (sucrose, glucose, fructose and starch) and nitrogenous compounds (nitrogen, total soluble proteins and free amino acids) were determined for leaves and developing seeds of P. vulgaris. Leaf tissue of elevated CO₂-grown plants accumulated significantly higher levels of both soluble sugars and starch. Leaf ultrastructure revealed considerable erilargement of starch grain sizes with surface areas more than five times larger compared to those of control plants. No apparent differences in structure and membrane integrity of chloroplasts in both CO₂ treatments were noted. Although ambient CO₂-grown plants had comparatively low levels of non-structural carbohydrates (NSC), they accumulated significantly higher levels of nitrogenous compounds. The levels of NSC were consistently higher in seeds of plants grown under elevated CO₂. In comparison to plants grown at elevated [CO₂], pods and seeds of ambient CO₂-grown plants had significantly larger pools of free amino compounds and N. Stomatal conductance (gs) declined significantly, as expected for plants grown under elevated CO₂. This was accompanied by a decline in transpiration rates (E). Reduced gs and E led to high AlE ratio, which meant improved water use efficiency (WUE) values for CO₂-enriched bean plants. Leaf carbon isotope discrimination (∆) against the heavier isotope of carbon (¹³C), has been used to select for high WUE in C₃ plants. In plants grown at elevated CO₂ concentration, ,1 was significantly reduced. Although ∆ was negatively correlated with WUE in both CO₂ treatments, the correlation was steeper and highly negative for CO₂-enriched plants. These results indicate underlying differences in gas-exchange physiology, including stomatal responses between ambient and elevated CO₂-grown plants. Photosynthetic acclimation was investigated using the response of assimilation to internal carbon dioxide concentration (A/C₁ curves). At early stages of growth, the initial slope of the A/C₁ response curve did not differ with CO₂ treatment. In contrast, CO₂-saturated photosynthetic rate (Amax) was significantly higher in plants grown under elevated versus ambient CO₂ at 15 DAG. However, at subsequent stages of growth both the initial slope and Amax declined in bean plants grown in elevated CO₂. Apparent carboxylation efficiency (ACE, estimated from the initial slope of A/C₁ response) values followed a similar trend and were significantly reduced in CO₂-enriched plants. These results indicate that acclimation or negative adjustment of photosynthesis may have been caused by a combination of both stomatal and biochemical limitations. Bean plants grown under conditions of elevated atmospheric CO₂ flowered 3 to 4 days earlier, and produced significantly more flowers and pods than plants grown at ambient conditions. Plants grown at elevated CO₂ aborted 22 and 20% more flowers and pods, respectively, than plants grown at ambient CO₂. Elevated CO₂ also significantly increased the number of tillers or lateral branches produced by plants, which contributed to a significant increase in pod number and seed yield in these plants. Although plants grown at elevated CO₂ produced on average 8 seeds per pod, while plants grown under ambient CO2 conditions produced 5 seeds per pod, the greater number of seeds was offset by lower seed weights in plants grown under _ elevated CO₂. Thus, despite high seed yield in beans grown under elevated CO₂, the harvest index (HI) did not change significantly between CO₂ treatments.
- Full Text:
- Date Issued: 1997
The oral-style South African short story in English A.W. Drayson to H.C. Bosman
- Authors: MacKenzie, Craig
- Date: 1997
- Subjects: Short stories, South African (English) -- History and criticism
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:2228 , http://hdl.handle.net/10962/d1002271
- Description: This study is concerned with a particular kind of short story in South African English literature - a kind of story variously called the fireside tale, tall tale, yarn, skaz narrative, frame narrative, or (the term used in this study), the 'oral-sty Ie story.' This kind of story is characterised by the use of an internal narrator (a fictional narrator or storyteller figure), the cadences of his or her speaking voice, and a 'reporting' frame narrator. Stories by A. W. Drayson, Frederick Boyle, J. Forsyth Ingram, W. C. Scully, Percy FitzPatrick, Ernest Glanville, Perceval Gibbon, Francis Carey Slater, Pauline Smith, Aegidius Jean Blignaut and Herman Charles Bosman form the principal body of primary sources examined in this study. The Bakhtinian notion of "simple" and "parodistic" skaz narratives is deployed to analyse the increasing complexity to be discerned in the works by these writers, which roughly span the 100 years from the middle of the nineteenth century to the middle of the present century. A "simple" use of the skaz narrative is evident in the early or 'ur-South African' oral-style story, represented here by Drayson, Boyle and Ingram. With Scully and FitzPatrick the form is still used 'artlessly,' although the beginnings of a greater self-consciousness can be discerned. The' Abe Pike' tales by Glanville introduce a more complex use of the fictional narrator, a process taken a step further by Gibbon in his 'Vrouw Grobelaar' tales. With the latter, in particular, the complex or "parodistic" skaz narrative makes its advent in South African literature. The oral-style stories of Slater and Smith are largely a regression to the ear lier form, although there are aspects of their stories which anticipate Bosman. With Blignaut and Bosman, however, the South African oral-style story comes into its own. In their Hottentot Ruiter and Oom Schalk Lourens characters is invested all the complexity and 'double-voicedness' that was latent, and largely dormant, in the earlier oral-style narratives. Through Blignaut, and Bosman in particular, the South African oral-style story achieves its most economical, sophisticated and successful form of expression. The study concludes by looking briefly at the use of an oral style in short stories by black South African writers and argues that their stories are not, formally speaking, to be categorised alongside those by the other~ writers examined. The oral-style story, the study concludes, achieved its apogee in Bosman's Oom Schalk Lourens sequence and went into sharp decline after Bosman's death in 1951.
- Full Text:
- Date Issued: 1997
- Authors: MacKenzie, Craig
- Date: 1997
- Subjects: Short stories, South African (English) -- History and criticism
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:2228 , http://hdl.handle.net/10962/d1002271
- Description: This study is concerned with a particular kind of short story in South African English literature - a kind of story variously called the fireside tale, tall tale, yarn, skaz narrative, frame narrative, or (the term used in this study), the 'oral-sty Ie story.' This kind of story is characterised by the use of an internal narrator (a fictional narrator or storyteller figure), the cadences of his or her speaking voice, and a 'reporting' frame narrator. Stories by A. W. Drayson, Frederick Boyle, J. Forsyth Ingram, W. C. Scully, Percy FitzPatrick, Ernest Glanville, Perceval Gibbon, Francis Carey Slater, Pauline Smith, Aegidius Jean Blignaut and Herman Charles Bosman form the principal body of primary sources examined in this study. The Bakhtinian notion of "simple" and "parodistic" skaz narratives is deployed to analyse the increasing complexity to be discerned in the works by these writers, which roughly span the 100 years from the middle of the nineteenth century to the middle of the present century. A "simple" use of the skaz narrative is evident in the early or 'ur-South African' oral-style story, represented here by Drayson, Boyle and Ingram. With Scully and FitzPatrick the form is still used 'artlessly,' although the beginnings of a greater self-consciousness can be discerned. The' Abe Pike' tales by Glanville introduce a more complex use of the fictional narrator, a process taken a step further by Gibbon in his 'Vrouw Grobelaar' tales. With the latter, in particular, the complex or "parodistic" skaz narrative makes its advent in South African literature. The oral-style stories of Slater and Smith are largely a regression to the ear lier form, although there are aspects of their stories which anticipate Bosman. With Blignaut and Bosman, however, the South African oral-style story comes into its own. In their Hottentot Ruiter and Oom Schalk Lourens characters is invested all the complexity and 'double-voicedness' that was latent, and largely dormant, in the earlier oral-style narratives. Through Blignaut, and Bosman in particular, the South African oral-style story achieves its most economical, sophisticated and successful form of expression. The study concludes by looking briefly at the use of an oral style in short stories by black South African writers and argues that their stories are not, formally speaking, to be categorised alongside those by the other~ writers examined. The oral-style story, the study concludes, achieved its apogee in Bosman's Oom Schalk Lourens sequence and went into sharp decline after Bosman's death in 1951.
- Full Text:
- Date Issued: 1997
Theories of exchange rates and the methodology of economics
- Authors: Hodge, Duncan
- Date: 1997
- Subjects: Foreign exchange rates Economics -- Methodology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:1012 , http://hdl.handle.net/10962/d1002747
- Description: This thesis is an exercise in applied methodology. Ideas in the history and philosophy of science which have proved to be influential in the methodology of economics, and in shaping economists' self-image in this regard, are selected for closer analysis and criticism. The main ideas that are addressed are those of empiricism, with emphasis on the methodological falsificationism of Karl Popper and Imre Lakatos, and Laudan's problem solving model of scientific progress . The thesis examines the relationship between empirical evidence, in the form of both econometric test results and stylized facts, and the development of theories about exchange rates and the open economy. This analysis begins with Cassel's formulation of purchasing power parity theory in 1916, through the elasticities, absorption, and Mundell-Fleming models of exchange rates and the balance of payments, up to the present day monetary and asset market models. This is done with regard to the broad methodological issues examined earlier in the thesis. Some of the main empirical and methodological difficulties in testing such theories are addressed, with particular reference to the role played by the Duhem-Quine thesis and the ceteris paribus assumption. Although some of these difficulties may be regarded as a matter of degree compared to similar problems in the natural sciences, it is argued that this difference is significant for the workability of falsification in economics . Moreover, the presence of hypotheses about expectations in many economic theories appears to be a substantive difference such that the difficulties posed by the Duhem-Quine thesis apply with much greater force in a social science like economics. The main conclusions are that neither the Popperian nor Lakatosian versions of falsification are seriously practiced in the area of exchange rate economics and that, unlike the position taken by advocates of falsification such as Mark Blaug, it would be inappropriate and misguided to do so. A tentative case is made, with some reference to the theories surveyed in this thesis, for the possibly greater relevance of Laudan' s more pragmatic problem solving model for the methodology of economics, particularly as regards his analysis and emphasis on conceptual problem solving in the progress of knowledge.
- Full Text:
- Date Issued: 1997
- Authors: Hodge, Duncan
- Date: 1997
- Subjects: Foreign exchange rates Economics -- Methodology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:1012 , http://hdl.handle.net/10962/d1002747
- Description: This thesis is an exercise in applied methodology. Ideas in the history and philosophy of science which have proved to be influential in the methodology of economics, and in shaping economists' self-image in this regard, are selected for closer analysis and criticism. The main ideas that are addressed are those of empiricism, with emphasis on the methodological falsificationism of Karl Popper and Imre Lakatos, and Laudan's problem solving model of scientific progress . The thesis examines the relationship between empirical evidence, in the form of both econometric test results and stylized facts, and the development of theories about exchange rates and the open economy. This analysis begins with Cassel's formulation of purchasing power parity theory in 1916, through the elasticities, absorption, and Mundell-Fleming models of exchange rates and the balance of payments, up to the present day monetary and asset market models. This is done with regard to the broad methodological issues examined earlier in the thesis. Some of the main empirical and methodological difficulties in testing such theories are addressed, with particular reference to the role played by the Duhem-Quine thesis and the ceteris paribus assumption. Although some of these difficulties may be regarded as a matter of degree compared to similar problems in the natural sciences, it is argued that this difference is significant for the workability of falsification in economics . Moreover, the presence of hypotheses about expectations in many economic theories appears to be a substantive difference such that the difficulties posed by the Duhem-Quine thesis apply with much greater force in a social science like economics. The main conclusions are that neither the Popperian nor Lakatosian versions of falsification are seriously practiced in the area of exchange rate economics and that, unlike the position taken by advocates of falsification such as Mark Blaug, it would be inappropriate and misguided to do so. A tentative case is made, with some reference to the theories surveyed in this thesis, for the possibly greater relevance of Laudan' s more pragmatic problem solving model for the methodology of economics, particularly as regards his analysis and emphasis on conceptual problem solving in the progress of knowledge.
- Full Text:
- Date Issued: 1997
Urban agriculture: advocacy and practice: a discursive study with particular reference to three Eastern Cape centres
- Authors: Webb, Nigel Leigh
- Date: 1997
- Subjects: Urban agriculture -- South Africa -- Eastern Cape Land use, Urban -- South Africa -- Eastern Cape Agriculture -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:2095 , http://hdl.handle.net/10962/d1002658
- Description: The purpose of this thesis is to explain the continued advocacy of 'urban agriculture' (UA) in the literature in the face of the seemingly modest role played by cultivation in practice. The analytical framework adopted isrthat of discourse, the theoretical underpinnings of which are derived from the early works of Foucault, and applications such as those of Escobar (1989; 1991) and Ferguson (1990). Using a discursive orientation involves two main tasks - an explanation of how 'UA' has gained some prominence and the man~er in which its currency is able to be maintained. The investigation included an in-depth analysis of the literature and empirical research in th!"e~Eastern Cape centres. The function of the empirical findings is to expose the discourse, as well as to extend the empirical base relating to 'urban agricultural' research in general. The thesis suggests that the growth in interest in 'UA' is a result of a Foucaultian "gap" opening in the discourse. Changes in the conceptualisation of development, the rise-in prominence of the urban poor and emerging ecodevelopment views, among others, have given proponents of "UA' greater room for manoeuvre. However, most case studies promote 'UA' despite providing little evidence of its role in household welfare. The way the discourse maintains its' currency is in the manner in which the objects of 'UA' are constituted. Firstly, the people are characterised as poor, largely divorced from their rural origins, and mainly female. Each of these assertions is questioned on the basis of the literature itself and the empirical findings. Secondly, in the practice of cultivation, emphasis is placed on the inputs used, and the crops that are produced. The treatment of both the inputs and the crops by the literature is cursory. Investigations show that land availability, garden size and irrigation water are e~pecially problematic and vegetable consumption remains low. Four types of alleged benefit associated with 'UA' are analysed - environmental and ecological, psychological and social, financial, and nutritional. In each case commonly held assumptions are undermined by closer investigation. Thus, the thesis contends that the role played by cultivation is a modest one. Most of the claims made on behalf of 'UA' are more congruent with development discourse than actual cultivation practice.
- Full Text:
- Date Issued: 1997
- Authors: Webb, Nigel Leigh
- Date: 1997
- Subjects: Urban agriculture -- South Africa -- Eastern Cape Land use, Urban -- South Africa -- Eastern Cape Agriculture -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:2095 , http://hdl.handle.net/10962/d1002658
- Description: The purpose of this thesis is to explain the continued advocacy of 'urban agriculture' (UA) in the literature in the face of the seemingly modest role played by cultivation in practice. The analytical framework adopted isrthat of discourse, the theoretical underpinnings of which are derived from the early works of Foucault, and applications such as those of Escobar (1989; 1991) and Ferguson (1990). Using a discursive orientation involves two main tasks - an explanation of how 'UA' has gained some prominence and the man~er in which its currency is able to be maintained. The investigation included an in-depth analysis of the literature and empirical research in th!"e~Eastern Cape centres. The function of the empirical findings is to expose the discourse, as well as to extend the empirical base relating to 'urban agricultural' research in general. The thesis suggests that the growth in interest in 'UA' is a result of a Foucaultian "gap" opening in the discourse. Changes in the conceptualisation of development, the rise-in prominence of the urban poor and emerging ecodevelopment views, among others, have given proponents of "UA' greater room for manoeuvre. However, most case studies promote 'UA' despite providing little evidence of its role in household welfare. The way the discourse maintains its' currency is in the manner in which the objects of 'UA' are constituted. Firstly, the people are characterised as poor, largely divorced from their rural origins, and mainly female. Each of these assertions is questioned on the basis of the literature itself and the empirical findings. Secondly, in the practice of cultivation, emphasis is placed on the inputs used, and the crops that are produced. The treatment of both the inputs and the crops by the literature is cursory. Investigations show that land availability, garden size and irrigation water are e~pecially problematic and vegetable consumption remains low. Four types of alleged benefit associated with 'UA' are analysed - environmental and ecological, psychological and social, financial, and nutritional. In each case commonly held assumptions are undermined by closer investigation. Thus, the thesis contends that the role played by cultivation is a modest one. Most of the claims made on behalf of 'UA' are more congruent with development discourse than actual cultivation practice.
- Full Text:
- Date Issued: 1997
Vitamin E supplementation and secondary metabolites interactions and effects on melanoma growth
- Authors: Ottino, Paulo
- Date: 1997
- Subjects: Vitamin E Melanoma
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4016 , http://hdl.handle.net/10962/d1004076
- Description: The present study was undertaken to determine the effects and possible mechanism of action of vitamin E succinate on malignant murine melanoma (BL6) and non-malignant monkey kidney (LLCMK) cell growth in vitro. Studies revealed that supplementation of 5, 7 and lOJLg/ml vitamin E succinate significantly inhibited BL6 cell growth, while in LLCMK cells no significant increase or decrease in growth was observed. The actual mechanism by which vitamin E succinate inhibits BL6 cell growth is at present unclear. Studies have suggested a radical or oxidant involvement in a number of degenerative diseases such as cancer, and that supplementation of antioxidant vitamins such as vitamin E may function to reduce cancer cell growth by quenching free radical species and preventing lipid peroxidation. In addition to its antioxidant role in a cell, vitamin E is believed to modulate the activities of various enzymes and metabolites in the eicosanoid pathway. Hence, this study investigated the effects of vitamin E succinate supplementation on free radical and lipid peroxidation levels, as well as the activities of various enzymes and metabolites ill the eicosanoid pathway. Throughout this study, emphasis was placed on BL6 melanoma cells since the magnitude of the relationship between LLCMK growth and the levels of various enzymes and metabolites in the eicosanoid pathway varied considerably from one experiment to another and did not show the consistent trend found with the BL6 cells. A decrease in cell growth was found to be accompanied by a concomitant increase rather than a decrease in the levels of free radicals and lipid peroxidation, suggesting that the growth inhibitory effects of vitamin E succinate on BL6 cells in vitro was not due to its antioxidant properties associated with the vitamin E component, but rather due to one or more of its other potential roles within the cell. This proposal was further strengthened by findings that vitamin E succinate, a non-physiological antioxidant in its esterified form, did not undergo significant cleavage to free vitamin E in the BL6 cells. Vitamin E succinate is believed to modulate membrane bound enzyme activities through physicochemical interactions with membrane lipids and changes in membrane fluidity. Hence, this study investigated the role of vitamin E succinate in modulating the activity of various enzymes and secondary messengers in the eicosanoid pathway. Supplementation of l-lOjLg/ml vitamin E succinate resulted in an overall increase in phospholipase A2 activity while cyclooxygenase and adenyl ate cyclase activities were found to be significantly increased at vitamin E succinate concentrations of 7 and WjLg/ml respectively. A significant increase in" 5-LOX activity was observed a! 10jLg/mi supplementation. The suggestion that vitamin E succinate modulates membrane bound enzyme activities was further strengthened by uptake and cellular distribution studies, which showed significantly higher levels of vitamin E succinate in membrane fractions of BL6 cells when compared with stroma fractions. Another factor which could account for elevated PLA2,-5-LOX and COX activities in BL6 cells as a result of vitamin E succinate supplementation, was that of intracellular calcium levels. Supplementation of BL6 cells with 1-7 jLg/ml vitamin E succinate resulted in an overall increase in intracellular calcium levels. These changes in calcium levels however were positively correlated with changes in PLA2 activity only. Since the rate of prostaglandin synthesis is controlled by phospholipase A2 activity, and net prostagiandin production is dependant on cyclooxygenase activity, the effects of vitamin E succinate supplementation on prostaglandin levels in BL6 cells was determined. Vitamin E succinate supplementation resulted in a significant decrease in prostaglandin D2 levels at vitamin E succinate concentrations of 3, 5, 7 and lOjLg/ml respectively, while prostaglandin F2a levels were significantly decreased at 1-10jLg/ml vitamin E succinate. The increases in prostaglandin E2 and 12 levels were inversely related to BL6 cell growth suggesting that both prostaglandins may act as negative regulators of BL6 cell growth. When comparing prostaglandin E2 levels to prostaglandin 12 levels in BL6 cells, significantly higher levels of prostaglandin E2 were found, suggesting that vitamin E succinate effects were mediated primarily through an increase in prostaglandin E2 levels. Furthermore, prostaglandin E2 levels are believed to modulate adenylate cyclase activity. It is therefore reasonable to conclude that the increased adenyl ate cyclase activity found in BL6 cells was dependant on prostaglandin E2 levels, since increases in prostaglandin E2 levels at 7 and lOjLg/ml vitamin E succinate correlated with an increase in adenylate cyclase activity and cyclic adenosine monophosphate levels. Thus it appeared that the observed inhibitory effects of vitamin E succinate supplementation on BL6 cell growth was not due to the antioxidant properties associated with the vitamin E component of the vitamin E succinate molecule, but was rather mediated in part through a cascade effect initiated by phospholipase A2 activation and archidonic acid release. This initial effect then appeared to result in an increase in cyclooxygenase activity and activation of a prostaglandin E2-adenylate cyclase-cyclic adenosine monophosphate linked system, ultimately altering cyclic adenosine monophosphate levels and inhibiting BL6 cell growth. This was confirmed when BL6 cells were supplemented with indomethacin, a cyclooxygenase inhibitor. Supplementation with the inhibitor resulted in vitamin E succinate having no inhibitory effects on BL6 cell growth. Furthermore, when comparing the levels of prostaglandin ~, adenylate cyclase activity and cyclIC adenosine monophosphate in the indomethacin treated cultures to non-indomethacin treated cultures, markedly lower levels of these metabolites were found in the indomethacin treated cultures. The cause of the increase in free radical and lipid peroxidation levels in BL6 cells following vitamin E succinate supplementation was further investigated. Cyclooxygenase enzymes are believed to generate free radical species and contribute to lipid peroxidation levels during catalytic activity. Markedly lower levels of free radicals and lipid peroxidation in indomethacin treated cultures were found when compared with vitamin E succinate treated cultures alone, suggesting that the increases in free radical and lipid peroxidation levels in BL6 cells supplemented with vitamin E succinate were indirectly due to an increase in cyclooxygenase activity in these cells.
- Full Text:
- Date Issued: 1997
- Authors: Ottino, Paulo
- Date: 1997
- Subjects: Vitamin E Melanoma
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4016 , http://hdl.handle.net/10962/d1004076
- Description: The present study was undertaken to determine the effects and possible mechanism of action of vitamin E succinate on malignant murine melanoma (BL6) and non-malignant monkey kidney (LLCMK) cell growth in vitro. Studies revealed that supplementation of 5, 7 and lOJLg/ml vitamin E succinate significantly inhibited BL6 cell growth, while in LLCMK cells no significant increase or decrease in growth was observed. The actual mechanism by which vitamin E succinate inhibits BL6 cell growth is at present unclear. Studies have suggested a radical or oxidant involvement in a number of degenerative diseases such as cancer, and that supplementation of antioxidant vitamins such as vitamin E may function to reduce cancer cell growth by quenching free radical species and preventing lipid peroxidation. In addition to its antioxidant role in a cell, vitamin E is believed to modulate the activities of various enzymes and metabolites in the eicosanoid pathway. Hence, this study investigated the effects of vitamin E succinate supplementation on free radical and lipid peroxidation levels, as well as the activities of various enzymes and metabolites ill the eicosanoid pathway. Throughout this study, emphasis was placed on BL6 melanoma cells since the magnitude of the relationship between LLCMK growth and the levels of various enzymes and metabolites in the eicosanoid pathway varied considerably from one experiment to another and did not show the consistent trend found with the BL6 cells. A decrease in cell growth was found to be accompanied by a concomitant increase rather than a decrease in the levels of free radicals and lipid peroxidation, suggesting that the growth inhibitory effects of vitamin E succinate on BL6 cells in vitro was not due to its antioxidant properties associated with the vitamin E component, but rather due to one or more of its other potential roles within the cell. This proposal was further strengthened by findings that vitamin E succinate, a non-physiological antioxidant in its esterified form, did not undergo significant cleavage to free vitamin E in the BL6 cells. Vitamin E succinate is believed to modulate membrane bound enzyme activities through physicochemical interactions with membrane lipids and changes in membrane fluidity. Hence, this study investigated the role of vitamin E succinate in modulating the activity of various enzymes and secondary messengers in the eicosanoid pathway. Supplementation of l-lOjLg/ml vitamin E succinate resulted in an overall increase in phospholipase A2 activity while cyclooxygenase and adenyl ate cyclase activities were found to be significantly increased at vitamin E succinate concentrations of 7 and WjLg/ml respectively. A significant increase in" 5-LOX activity was observed a! 10jLg/mi supplementation. The suggestion that vitamin E succinate modulates membrane bound enzyme activities was further strengthened by uptake and cellular distribution studies, which showed significantly higher levels of vitamin E succinate in membrane fractions of BL6 cells when compared with stroma fractions. Another factor which could account for elevated PLA2,-5-LOX and COX activities in BL6 cells as a result of vitamin E succinate supplementation, was that of intracellular calcium levels. Supplementation of BL6 cells with 1-7 jLg/ml vitamin E succinate resulted in an overall increase in intracellular calcium levels. These changes in calcium levels however were positively correlated with changes in PLA2 activity only. Since the rate of prostaglandin synthesis is controlled by phospholipase A2 activity, and net prostagiandin production is dependant on cyclooxygenase activity, the effects of vitamin E succinate supplementation on prostaglandin levels in BL6 cells was determined. Vitamin E succinate supplementation resulted in a significant decrease in prostaglandin D2 levels at vitamin E succinate concentrations of 3, 5, 7 and lOjLg/ml respectively, while prostaglandin F2a levels were significantly decreased at 1-10jLg/ml vitamin E succinate. The increases in prostaglandin E2 and 12 levels were inversely related to BL6 cell growth suggesting that both prostaglandins may act as negative regulators of BL6 cell growth. When comparing prostaglandin E2 levels to prostaglandin 12 levels in BL6 cells, significantly higher levels of prostaglandin E2 were found, suggesting that vitamin E succinate effects were mediated primarily through an increase in prostaglandin E2 levels. Furthermore, prostaglandin E2 levels are believed to modulate adenylate cyclase activity. It is therefore reasonable to conclude that the increased adenyl ate cyclase activity found in BL6 cells was dependant on prostaglandin E2 levels, since increases in prostaglandin E2 levels at 7 and lOjLg/ml vitamin E succinate correlated with an increase in adenylate cyclase activity and cyclic adenosine monophosphate levels. Thus it appeared that the observed inhibitory effects of vitamin E succinate supplementation on BL6 cell growth was not due to the antioxidant properties associated with the vitamin E component of the vitamin E succinate molecule, but was rather mediated in part through a cascade effect initiated by phospholipase A2 activation and archidonic acid release. This initial effect then appeared to result in an increase in cyclooxygenase activity and activation of a prostaglandin E2-adenylate cyclase-cyclic adenosine monophosphate linked system, ultimately altering cyclic adenosine monophosphate levels and inhibiting BL6 cell growth. This was confirmed when BL6 cells were supplemented with indomethacin, a cyclooxygenase inhibitor. Supplementation with the inhibitor resulted in vitamin E succinate having no inhibitory effects on BL6 cell growth. Furthermore, when comparing the levels of prostaglandin ~, adenylate cyclase activity and cyclIC adenosine monophosphate in the indomethacin treated cultures to non-indomethacin treated cultures, markedly lower levels of these metabolites were found in the indomethacin treated cultures. The cause of the increase in free radical and lipid peroxidation levels in BL6 cells following vitamin E succinate supplementation was further investigated. Cyclooxygenase enzymes are believed to generate free radical species and contribute to lipid peroxidation levels during catalytic activity. Markedly lower levels of free radicals and lipid peroxidation in indomethacin treated cultures were found when compared with vitamin E succinate treated cultures alone, suggesting that the increases in free radical and lipid peroxidation levels in BL6 cells supplemented with vitamin E succinate were indirectly due to an increase in cyclooxygenase activity in these cells.
- Full Text:
- Date Issued: 1997