The sacred lotus (Nelumbo nucifera) is a large aquatic plant endemic to subtropical and tropical Asia and northern Australia. Lotus has a combination of morphological and anatomical features that make it challenging for research work. The necessity of research on lotus is driven by niche market opportunities identified in Japan during counter seasonal production periods. Several features of lotus are utilised for consumer applications with commercial promise including seeds, young shoots and rhizome production. Further, the flowers and seed pods have value as cut-flower products and religious decoration. Several challenges have to be overcome before production of adequate products can be realised in Australia. The challenges which can be addressed most immediately are the questions regarding plant nutrition for lotus. In order to accomplish certain objectives, development of a system for growth and analysis of imposed nutrient treatments in replication, which accommodated a plant with unusual and seasonal attributes, was essential. The results of the tests conducted on the trialled system, whilst not conclusive, provide a solid reference for any future research on lotus nutrition. Recommendations are made for design and enhancement of the system to provide guidelines for such research. / Doctor of Philosophy (PhD)

Compared with the free atmosphere, the aquatic environment is oxygen poor. As a result many secondarily aquatic organisms have adaptations that allow them to continue to use the atmosphere, directly or indirectly, to supply their oxygen requirements. This thesis examines how diving insects use bubbles of air collected at the surface of the water as oxygen reserves, gills and flotation devices, and how an aquatic angiosperm channels convective flows of air from its emergent leaves to its submerged organs. 1. Backswimmers (Anisops spp.) begin a dive positively buoyant, but rapidly enter a protracted period of near neutral buoyancy. A bubble of air held on the insect’s abdomen shrinks as respiration consumes its oxygen, while at the same time highly soluble carbon dioxide dissolves into the surrounding water. The reduced air volume confers neutral buoyancy. In response to low oxygen partial pressure (PO2) in the bubble, oxygen is released from large haemoglobin cells in the abdomen. The haemoglobin’s sensitivity to falling PO2 maintains the oxygen tension between 5.1 and 2.0 kPa. This stabilises the volume and buoyancy of the bubble. During a dive the haemoglobin and air-store supply 0.25 and 0.26 μL of oxygen, respectively. 2. The oxygen affinity of backswimmer haemoglobin determines the stability of the neutrally buoyant phase as well as its ability to satisfy the insect’s respiration. An oxygen equilibrium curve (OEC) determined in vivo has a highly sigmoid shape and an oxygen affinity of 3.9 kPa. In comparison with OEC made in vitro, the in vivo measurements show increased cooperativity and oxygen affinity, consistent with the presence of cationic effectors. Models strongly support the accuracy of the in vivo OEC method. 3. It has long been assumed that a bubble of air held over the spiracles of an insect enables the uptake of oxygen from the surrounding water and thus acts as a ‘gas gill’. Oxygen diffuses into a bubble of air when its PO2 is lower than the surrounding water, but a coincident higher nitrogen partial pressure causes it to dissolve. Several models have been produced to describe the gas exchange process, but all are based on untested assumptions of gill parameters. Measurements of gas gill volume and PO2 made on water bugs (Agraptocorixa eurynome) demonstrate that both drop quickly at the beginning of a dive, but PO2 reaches a stable level while the gas gill continues to dissolve. The importance of ventilation in maintaining an acceptable rate of oxygen consumption is also shown. 4. Interconnected gas spaces within the leaves, stems and rhizomes are a common feature of many emergent aquatic plants. Pressurised air from the leaves and culms of these plants ventilate these lacunae, flowing back to the atmosphere through efflux points. Unlike most aquatic plants, which have simple interconnected pith spaces, sacred lotus, Nelumbo nucifera, possess discrete gas canals which only interconnect where a leaf grows from the rhizome. Silicone casts and pneumatic tests of the gas canals reveal a complex repeating pattern of interconnections which channel air from specific regions of the leaf blade to the rhizome and out through efflux points on adjacent leaves. 5. Lotus, Nelumbo nucifera, possess in the centre of their leaves a specialised efflux organ which connects the gas canals in the leaves and stems with the atmosphere through the apertures of large stomata. Measurements made on excised lotus leaves and in situ reveal that the large stomata act as exhaust valves, opening and closing in a diurnal pattern to regulate the flow of pressurised gas from the leaf lamina and gas canals. This behaviour is shown to regulate gas flow rate and direction. The aquatic environment offers similar respiratory challenges to both plants and insects. While the oxygen uptake and transport mechanisms evolved by these groups are markedly different, they all function according to the same physical laws. Diving insects are separated from the atmosphere while underwater, forcing them to rely on oxygen either carried with them from the surface or extracted from the surrounding water. Emergent aquatic plants have permanent access to atmospheric oxygen, but must transport it long distances from their aerial leaves and stems to their roots and rhizomes. This thesis examines the uptake and storage of oxygen by diving insects and the gas transport system of the sacred lotus. / Thesis(Ph.D.) -- University of Adelaide, School of Earth and Environmental Sciences, 2008