Graduation date: 1997

Ecophysiological responses of Litopenaeus vannamei were evaluated as functions of 1) salinity and animal size, 2) temperature and the animal's nutritive state, and 3) dissolved-oxygen concentration and animal size. Growth rate, routine metabolic rate, limiting oxygen concentration for routine metabolism, and marginal metabolic scope were determined for L. vannamei maintained and tested at salinities of 2, 10, and 28 ppt, all at 28 C. Routine metabolic rate (RMR) was not demonstrably dependent on salinity but decreased with increasing shrimp weight. Limiting oxygen concentration for routine metabolism (LOCr) was independent of shrimp weight up to 9 g; but, for larger shrimp, decreased with increasing weight. Marginal metabolic scope (MMS = RMR/LOCr) also decreased with increasing shrimp weight and was independent of salinity for shrimp weighing up to 9 g; but, like LOCr, MMS was dependent on salinity for larger shrimp. Growth rate was significantly less at 2 ppt than at 10 or 28 ppt, which gave similar growth rates. The effects of four temperatures (20, 24, 28, and 32 C) on growth, RMR, LOCr, and MMS were examined for fed and starved L. vannamei. Routine metabolic rate increased with increased temperature both for fed and starved shrimp. Marginal metabolic scope and growth appeared to be positively related and, at 20 C, seemed to induce a state of metabolic torpor. Data from the study of chronic effects of hypoxia (~2 mg O2 L-1) vs. normoxia (> 5 mg O2 L-1) on ecophysiological responses indicated that although low-DO environments can depress RMR and growth in L. vannamei, animals grown under hypoxic and normoxic conditions did not differ in their metabolic responses upon acute exposure to hypoxia, providing no evidence of acclimation to hypoxia in L. vannamei. Data from the above experiments were used to parameterize Ecophys.Shrimp, a computer simulation model of shrimp growth in time-varying environmental regimes. One unified model was able to simulate all my experiments; and, with only minimal adjustment of the model parameter MMSO, it also adequately simulated studies taken from the literature. Thus, Ecophys.Shrimp seems capable of realistically representing the ecophysiological dynamics of shrimp metabolism and growth in various culture systems.

The C3 photosynthetic pathway uses light energy to reduce CO2 to carbohydrates and other organic compounds and is a central component of biological metabolism. In C3 photosynthesis, CO2 assimilation is catalyzed by ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), which reacts with both CO2 and O2. While competitive inhibition of CO2 assimilation by oxygen is suppressed at high CO2 concentrations, O2 inhibition is substantial when CO2 concentration is low and O2 concentration is high; this inhibition is amplified by high temperature and aridity (Sage 2004). Atmospheric CO2 concentration dropped below saturating levels 25-30 million years ago (Tipple & Pagani 2007), and the C4 photosynthetic pathway is hypothesized to have first evolved in warm, low latitude environments around this time (Christin et al. 2008a). The primary feature of C4 photosynthesis is suppression of O2 inhibition through concentration of CO2 around Rubisco. This pathway is estimated to have evolved almost 50 times across 19 angiosperm families (Muhaidat et al. 2007), a remarkable example of evolutionary convergence. In several C4 lineages, there are species with photosynthetic traits that are intermediate between the C3 and C4 states, known as C3-C4 intermediates. In two eudicot genera, Flaveria (Asteraceae) and Alternanthera (Amaranthaceae), there is evidence that these species represented an intermediate state in the evolution of the C4 pathway (McKown et al. 2005; Sanchez-del Pino 2009). The purpose of this thesis is to ascertain the specific benefits to plant carbon balance and resource-use efficiencies of the C3-C4 pathway relative to C3 species, particularly at low CO2 concentrations and high temperatures, factors which are thought to have been important in selecting for C3-C4 traits (Ehleringer et al. 1991). This will provide information on the particular advantages of the C3-C4 pathway in warm, often arid environments and how these advantages may have been important in advancing the initial stages of C4 evolution in eudicots. This thesis addresses the physiological intermediacy of previously uncharacterized C3-C4 species of Heliotropium (Boraginaceae); the water- and nitrogen-use efficiencies of C3-C4 species of Flaveria; and the photosynthetic performance and acclimation of C3, C4 and C3-C4 species of Heliotropium, Flaveria and Alternanthera grown at low and current ambient CO2 levels and high temperature.

The St. Lucia Estuary, Africa’s largest estuarine lake, is currently experiencing an unprecedented crisis related to freshwater deprivation. This has resulted in a reversed salinity gradient and drastically reduced water levels. These harsh environmental conditions, combined with the limited connection with the open ocean have lead to a loss of biodiversity in the system. The dominant zooplankton taxa include the copepods Pseudodiaptomus stuhlmanni and Acartia natalensis and the mysid Mesopodopsis africana. In March 2007, the closed-mouth state was briefly interrupted by an open-mouth phase, induced by a unique combination of extreme climatic events. With the incoming seawater, previously excluded marine taxa re-entered the system, increasing its diversity significantly. Salinity and temperature have been referred to as driving forces in aquatic ecosystems. The tolerance limits of the key mysid species were, therefore, investigated. Results showed that M. africana has some of the highest recorded upper salinity and temperature tolerances for a mysid. Because of their high biomass, mysids have the potential to affect microalgal standing stocks. Their grazing dynamics (in relation to autotrophic food availability) were investigated in two contrasting environments within the estuary. Ingestion rates and subsequently population grazing impacts on the total microalgal standing stocks were higher at the Mouth than at Charters Creek. This was attributed to the harsh environmental conditions in the latter region. Despite the lower ingestion rates exhibited here, these mysids seem capable of meeting their energetic requirements from a microalgal diet alone. Stable isotope data, though, show that they also utilise a heterotrophic diet. Results of the mixed model SIAR v 4 revealed the contribution of the different carbon sources to the diet of M. africana. Most unique was this mysid’s ability to modify its diet on both short temporal and spatial scales. Resource utilization between the dominant taxa was also compared. All three taxa appear to be opportunistic feeders, capable of incorporating a number of food items in their diet. Between food partitioning, predator avoidance strategies, and their common ability to survive in highly dynamic environments, these species are capable of co-existing, and together contribute to the overall resilience so far shown by the system. / Thesis (Ph.D.)-University of KwaZulu-Natal, Westville, 2011.

The C3 photosynthetic pathway uses light energy to reduce CO2 to carbohydrates and other organic compounds and is a central component of biological metabolism. In C3 photosynthesis, CO2 assimilation is catalyzed by ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), which reacts with both CO2 and O2. While competitive inhibition of CO2 assimilation by oxygen is suppressed at high CO2 concentrations, O2 inhibition is substantial when CO2 concentration is low and O2 concentration is high; this inhibition is amplified by high temperature and aridity (Sage 2004). Atmospheric CO2 concentration dropped below saturating levels 25-30 million years ago (Tipple & Pagani 2007), and the C4 photosynthetic pathway is hypothesized to have first evolved in warm, low latitude environments around this time (Christin et al. 2008a). The primary feature of C4 photosynthesis is suppression of O2 inhibition through concentration of CO2 around Rubisco. This pathway is estimated to have evolved almost 50 times across 19 angiosperm families (Muhaidat et al. 2007), a remarkable example of evolutionary convergence. In several C4 lineages, there are species with photosynthetic traits that are intermediate between the C3 and C4 states, known as C3-C4 intermediates. In two eudicot genera, Flaveria (Asteraceae) and Alternanthera (Amaranthaceae), there is evidence that these species represented an intermediate state in the evolution of the C4 pathway (McKown et al. 2005; Sanchez-del Pino 2009). The purpose of this thesis is to ascertain the specific benefits to plant carbon balance and resource-use efficiencies of the C3-C4 pathway relative to C3 species, particularly at low CO2 concentrations and high temperatures, factors which are thought to have been important in selecting for C3-C4 traits (Ehleringer et al. 1991). This will provide information on the particular advantages of the C3-C4 pathway in warm, often arid environments and how these advantages may have been important in advancing the initial stages of C4 evolution in eudicots. This thesis addresses the physiological intermediacy of previously uncharacterized C3-C4 species of Heliotropium (Boraginaceae); the water- and nitrogen-use efficiencies of C3-C4 species of Flaveria; and the photosynthetic performance and acclimation of C3, C4 and C3-C4 species of Heliotropium, Flaveria and Alternanthera grown at low and current ambient CO2 levels and high temperature.

Seaweeds that occupy the intertidal zone grow in distinct vertical bands. Elucidating the causes of zonation of intertidal seaweeds has been a key question for over a century, and the prevailing paradigm is that species occupying the higher shore position are limited by abiotic factors associated with exposure to atmospheric conditions, whereas for the lower-shore species competition is considered to limit a macroalga�s distribution. The main objective of this study was to assess the effect of desiccation, and to a lesser extent nutrient limitation, on the distribution of Stictosiphonia arbuscula, a red intertidal seaweed that grows in a wide vertical band in the mid - high shore region. A multi-disciplinary approach was used to investigate the effect of desiccation at different functional levels, from individual cells to the whole organism. Ecophysiological comparisons were made on high-shore and low-shore populations to determine whether the upper and lower shore distributions have different abilities to tolerate and recover from desiccation stress during summer and winter. The effect of desiccation was visually assessed, using electron microscopy, at the cellular and whole organism level. Carbohydrate and pigment concentrations were determined to provide information on how S. arbuscula allocates its resources between winter and summer. Investigations into the ability of Stictosiphonia arbuscula to tolerate and recover photosynthetic capacity after desiccation events (relative humidity (R.H.) 5% and 40%; ranging from 30 minutes - 24 hours) unexpectedly revealed no differences between high- and low-shore specimens. However, there were seasonal differences, with summer specimens better able to recover both tissue water content and photosynthetic capacity after desiccation compared to winter specimens, especially after desiccation for 24 hours. In contrast high-shore specimens had increased rates of nitrogen uptake after mild desiccation treatments, compared to low-shore specimens during summer, and specimens from the low-shore had increased variability in their uptake rates, indicating disruption to the uptake mechanisms, compared to the high-shore specimens. S. arbuscula was able to recover photosynthetic capacity faster than nitrogen and phosphorus uptake during re-immersion after desiccation, which indicates that the ability to regain photosynthetic function may be more important for this species than regaining the ability to take up nitrogen and phosphorus. As S. arbuscula can photosynthesise in air (providing there is sufficient tissue water content) and water, the �window of opportunity� to assimilate carbon is greater than for nitrogen and phosphorus uptake, which is only available to S. arbuscula during immersion at high tides. Despite this, no evidence was found of severe nutrient limitation in either high- or low-shore specimens. The disruption and recovery of cellular organisation during rehydration, assessed visually using electron microscopy, showed that the rate and duration of desiccation also affected the extent of recovery. Extremely desiccating conditions (R.H. 5%) damaged the cells and no recovery was observed during rehydration, compared to moderate desiccation treatments (R.H. 40%) where the time taken to recover during rehydration increased with increasing desiccation periods. The disruption of cellular organisation observed in Stictosiphonia arbuscula cells, was found to match the reduction and / or disruption to photosynthesis and the uptake of nitrogen and phosphorus under similar desiccation regimes. The formation of ridges and depressions on the surface of the tissue was also observed during desiccation, which is thought to help reduce the rate of evaporation by trapping �pockets� of air. Finally, Stictosiphonia arbuscula is extremely well suited to the mid - high region in the intertidal zone and it can maximise its competitive ability within this niche during immersion by fully rehydrating within 10 minutes, regaining maximum photosynthetic capacity within 20 minutes, and stability of nutrient uptake rates within 90 minutes. Further, S. arbuscula has higher rates of photosynthesis in winter, associated with increased photosynthetic pigment concentrations at this time. In contrast, during summer, the photosynthetic rates, chlorophyll a and phycobiliprotein concentrations are reduced and S. arbuscula increases its allocation of resources into protective mechanisms such as UV-absorbing compounds.

Ecophysiological attributes could be causes for rarity in plants. We tested the hypothesis that a species’ ability to regulate photosynthesis and growth in response to environmental factors is indicative of its environmental resilience and that this is linked to its conservation status. In this study, the ecophysiology of Graptophyllum reticulatum, an Australian endangered endemic species, was compared with that of its three closely related and more common congeners G. ilicifolium, G. excelsum and G. spinigerum. Ecophysiological attributes were measured on the four species in their natural habitats and under artificially imposed environmental stresses, including changed soil conditions, excess light and low water availability, in a glasshouse experiment. Photosynthesis was determined at the photosystem II and leaf level using chlorophyll a fluorescence and gas exchange techniques. Applied to the chlorophyll fluorescence transient of leaves, the JIP test provides a Performance Index which quantifies the main steps in PSII photochemistry including light energy absorption, excitation energy trapping, and conversion of excitation energy into electron flow. At the leaf level, gas exchange measurements allow determination of maximum CO2 assimilation rates, intercellular CO2 concentrations, stomatal conductance for water vapour and instantaneous water use efficiency. Growth analysis was performed to assess relative growth rates and physiological and morphological responses. Analysis of physiological differences and responses indicated that, compared to its more common relatives, the endangered G. reticulatum was an intrinsically slow growing species, exhibited the lowest fitness when growing in favorable environments and was most sensitive to excess light stress. Photoinhibition is therefore likely to restrict the endangered species to shade habitats. Compared with the endangered G. reticulatum, the vulnerable G. ilicifolium and common G. spinigerum species were better adapted to high light and changed nutrient levels, but were more susceptible to water stress. The rare G. excelsum had the fastest growth rate and the highest fitness in favorable environments. Based on the ecophysiological attributes examined here, it is proposed that excess light is likely to be the most critical abiotic factor restricting distribution of the endangered species in a fragmented landscape. The survival of the species may be most dependent on the intactness of the habitat over-storey. In contrast, the vulnerable G. ilicifolium showed strong susceptibility to water limitation, and survival might be threatened if climate change alters habitat water relations to cause, for example, more pronounced dry periods. The rare G. excelsum which had highest carbon gain and growth in the experiments carried out in this study, may become the most successful adaptation out of the rainforest environment due to its tolerance to higher light and limited water availability. To examine the generality of the link between rarity and ecophysiology with Graptophyllum species, two dipterocarp species, narrowly endemic Dipterocarpus condorensis and local common Shorea roxburghii that are actually co-located in South-eastern Vietnam were studied. Findings in this case study confirmed the usefulness of the comparative approach based on physiological measurements, either in situ or ex situ, to explain plant rarity. The results of this study indicate ecophysiological research is a tool for examining causes of rarity and possible abiotic threats. The information gained allows assessment of environmental resilience of species and contributes essential knowledge for management and conservation of threatened plants. Such knowledge is also useful for ex situ conservation including propagation, translocation and re-introduction in restoration programs.

Bibliography: leaves 148-161. / 168 leaves : ill. (some col.) ; 30 cm. / Title page, contents and abstract only. The complete thesis in print form is available from the University Library. / Thesis (Ph.D.)--Adelaide University, Dept. of Environmental Biology, 2001

Ecophysiological attributes could be causes for rarity in plants. We tested the hypothesis that a species’ ability to regulate photosynthesis and growth in response to environmental factors is indicative of its environmental resilience and that this is linked to its conservation status. In this study, the ecophysiology of Graptophyllum reticulatum, an Australian endangered endemic species, was compared with that of its three closely related and more common congeners G. ilicifolium, G. excelsum and G. spinigerum. Ecophysiological attributes were measured on the four species in their natural habitats and under artificially imposed environmental stresses, including changed soil conditions, excess light and low water availability, in a glasshouse experiment. Photosynthesis was determined at the photosystem II and leaf level using chlorophyll a fluorescence and gas exchange techniques. Applied to the chlorophyll fluorescence transient of leaves, the JIP test provides a Performance Index which quantifies the main steps in PSII photochemistry including light energy absorption, excitation energy trapping, and conversion of excitation energy into electron flow. At the leaf level, gas exchange measurements allow determination of maximum CO2 assimilation rates, intercellular CO2 concentrations, stomatal conductance for water vapour and instantaneous water use efficiency. Growth analysis was performed to assess relative growth rates and physiological and morphological responses. Analysis of physiological differences and responses indicated that, compared to its more common relatives, the endangered G. reticulatum was an intrinsically slow growing species, exhibited the lowest fitness when growing in favorable environments and was most sensitive to excess light stress. Photoinhibition is therefore likely to restrict the endangered species to shade habitats. Compared with the endangered G. reticulatum, the vulnerable G. ilicifolium and common G. spinigerum species were better adapted to high light and changed nutrient levels, but were more susceptible to water stress. The rare G. excelsum had the fastest growth rate and the highest fitness in favorable environments. Based on the ecophysiological attributes examined here, it is proposed that excess light is likely to be the most critical abiotic factor restricting distribution of the endangered species in a fragmented landscape. The survival of the species may be most dependent on the intactness of the habitat over-storey. In contrast, the vulnerable G. ilicifolium showed strong susceptibility to water limitation, and survival might be threatened if climate change alters habitat water relations to cause, for example, more pronounced dry periods. The rare G. excelsum which had highest carbon gain and growth in the experiments carried out in this study, may become the most successful adaptation out of the rainforest environment due to its tolerance to higher light and limited water availability. To examine the generality of the link between rarity and ecophysiology with Graptophyllum species, two dipterocarp species, narrowly endemic Dipterocarpus condorensis and local common Shorea roxburghii that are actually co-located in South-eastern Vietnam were studied. Findings in this case study confirmed the usefulness of the comparative approach based on physiological measurements, either in situ or ex situ, to explain plant rarity. The results of this study indicate ecophysiological research is a tool for examining causes of rarity and possible abiotic threats. The information gained allows assessment of environmental resilience of species and contributes essential knowledge for management and conservation of threatened plants. Such knowledge is also useful for ex situ conservation including propagation, translocation and re-introduction in restoration programs.

"June 2002" / Bibliography: leaves 172-198. / viii, 198 leaves : ill. ; 30 cm. / Title page, contents and abstract only. The complete thesis in print form is available from the University Library. / Proposes that the overall outcome of plant interactions along a temporal gradient of resource availability changes from positive during interpulses to negative during pulses. Examines negative interactions between 4 co-dominant chenopod scrubs in arid Acacia papyrocarpa woodlands. Negative interactions were more intense when conditions were least productive. Positive interactions between seedlings also changed over time, depending on the facilitation mechanism. Plant interactions seem to be most intense at the beginning of interpulses when plants are competing for diminishing water, or survivorship is enhanced in the favorable microsites provided by other plants. Later in the interpulse, interactions become less intense as conditions become more stressful and therefore survivorship and growth are affected more by abiotic conditions than plant interactions. / Thesis (Ph.D.)--University of Adelaide, Dept. of Environmental Biology, 2002