School of Agriculture, Food and Ecosystem Sciences - Theses
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ItemPhotosynthetic responses to light, nitrogen, phosphorus and pruning of Eucalyptus in south-eastern Australia( 2005)Eucalypts frequently grow faster after additions of fertiliser, but more slowly in the shade or following `green pruning'. The coupling of rates of growth to environmental factors is at least partly due to acclimation of photosynthetic processes. Photosynthesis rarely proceeds at maximum rates in natural environments as photosynthetic processes and the supply of basic requirements of photosynthesis (CO2, H20, light, phosphorus and nitrogen) vary at both short (minutes to hours) and longer (days to months) time scales. Currently we lack mechanistic explanations for how these variables, alone and in combination underpin changed growth rates in Eucalyptus. This study examined growth and photosynthetic characteristics in glasshouse-grown seedlings and field-grown trees of Eucalyptus species that are commonly planted for forestry and revegetation purposes in central Victoria. Acclimation to light (among seedlings and within canopies), nutrient availability (phosphorus and nitrogen) and increased sink-strength for photosynthates were the primary foci of the study. In each instance I examined distribution of leaf nutrients within a canopy and allocation of N to Rubisco and chlorophyll to assess the degree to which nutrients limit photosynthesis in Eucalyptus. A novel technique was introduced to quantify the allocation of inorganic phosphorus within cells (cytoplasm versus vacuole), followed by an assessment of inorganic phosphorus allocation in response to a long-term reduction in phosphorus supply. In all circumstances, rates of growth were responsive to environmental conditions. Growth responses were underpinned by altered patterns of biomass partitioning and changed leaf morphology more than by rates of photosynthesis per se. There was little difference in adaptive strategies implemented by seedlings and trees: both were oriented towards the accumulation of nutrients rather than increasing rates of photosynthesis. Photosynthesis was reduced by shading (among different plants and within the canopy of a tree) and reduced phosphorus supply whereas N had little effect on photosynthesis. Analysis of pools of inorganic P revealed that adequate supplies were maintained for photosynthetic processes regardless of P supply, therefore reduced photosynthesis follows, rather than leads, a more general leaf-level response to reduced P. Similarly, changed partitioning of nitrogen between Rubisco and chlorophyll was unnecessary as leaf nitrogen concentrations were consistently maintained at well above published minimum levels. Hence, photosynthesis was not up-regulated following increased nitrogen or phosphorus supply; instead excess nutrients were accumulated and used to support increased biomass. One exception was after defoliation, when up-regulation of photosynthesis was observed, presumably to ensure the demand for photosynthates could be met by a reduced leaf area. Sensitivity analyses consistently revealed variation in photosynthetic rates owed more to altered biochemical activity (e.g. Jmax and Vcmax) rather than stomatal conductance regardless of growth condition (glasshouse versus field). Hence, whilst Eucalyptus has considerable photosynthetic potential, faster rates of carbon fixation are only exhibited in the short-term. In part, this is due to the multiplicity of factors involved in `optimisation' of photosynthesis and their individual and collective responses to environmental conditions. In the long term however, increased canopy photosynthetic capacity follows only an increased photosynthetic area.