|dc.description.abstract||Background: This study seeks to model aspects of the regeneration of radiata pine (Pinus radiata D.Don) seedlings under a range of environmental conditions. This study investigated whether “hybrid” mechanistic models, which predict plant growth and development using empirical representations of plant physiological responses to the environment, could provide a realistic alternative to conventional empirical regeneration models.
Objectives: The objectives of this study were to 1) identify the functional relationships between the environmental conditions controlling germination, establishment and growth of radiata pine seedlings, under a range of those environmental conditions as specified by temperature and available light and soil water; and 2) specify those functional relationships in hybrid mechanistic (“hybrid”) models.
Methods: Radiata pine seedling germination and growth were measured under controlled environmental conditions (incubators for seed germination, growth cabinets for seedlings), and results used to adapt, parameterise and test two published hybrid models; one for germination (the hydrothermal time model); and one for seedling growth in the first six months after germination, based on plant radiation use efficiency (RUE).
The hydrothermal model was tested by incubating commercial radiata pine seeds under factorial combinations of temperature and water potentials where germination was likely to occur (12.5 ºC to 32.5 ºC and 0 MPa to –1.2 MPa.). 100 seeds were germinated for each factorial combination. The hydrothermal germination model was fitted to the germination data using non-linear regression modles, will allowed simultaneous estimation of all modle parameters.
Seedlings were grown in controlled growth cabinets, and their RUE was calculated as the ratio of net primary production (NPP, specified in terms of an increase in oven dry biomass), to PAR intercepted or absorbed by a seedling. Estimation of seedling RUE required development of novel techniques for non-destructive estimation of seedling oven dry weight, and measurement of PAR interception by seedlings. The effect of varying PAR flux density on RUE was tested by measuring RUE of seedlings grown at 125, 250 and 500 µmol m⁻² s⁻¹. In a second experiment, the effect of deficits in available soil water on RUE was tested by measuring RUE of seedlings grown under 250 µmol m⁻² s⁻¹ PAR flux, and at different levels of available soil water. Available soil water was specified by a soil moisture modifier factor (ƒθ) which ranges between 1 for moist soils and 0 for soils where there is insufficient water for seedling growth. This soil moisture modifier had not previously been applied in studies of tree seedling growth.
Temperatures for both seedling experiments were a constant 17.5 ºC (day) and 12.5 ºC (night).
Results: Hydrothermal time models accurately described radiata pine seed germination. Model predictions were closely correlated with actual seed germination over the full range of temperature and water potentials where germination was likely to occur (12.5 ºC to 32.5 ºC and 0 MPa to –1.2 MPa. The minimum temperature for germination (base temperature) was 9.0 ºC. Optimum temperatures for germination ranged from ~20ºC for slow-germinating seeds to ~27 ºC for the fastest germinating seeds. The minimum water potential for seed germination varied within the seed population, with an approximately normal distribution (base water potential = –1.38 MPa, standard deviation of 0.48 MPa). In the process of developing the model, a novel explanation for the decline in germination rates at supra-optimal temperatures was developed (Section 3.4.6), based on earlier models proposed by Alvarado & Bradford (2002) and Rowse & Finch-Savage (2003). This explanation was that the decline in germination rate was not driven just by temperature, but by accumulated hydrothermal time above the base temperature for germination (T₀). This in turn raised the base soil water potential (Ψb) towards 0, so that the reduction in germination rate arose from a reduced accumulation of hydro-time, rather than from thermal denaturation of enzymes facilitating germination – the conventional explanation for non-linear accumulation of thermal time at supra-optimal temperatures for plant development. Upwards adjustment (towards 0 MPa) of base water potentials of germinating seeds occurred also at very cold temperatures in combination with high water potentials. In both cases (very cold or else supra-optimal temperatures) this upwards adjustment in base water potentials prevented germination of part of the seed population, and is proposed as a mechanism which enables seed populations to “hedge their bets” when germinating under less than ideal germination conditions.
RUE of young germinated radiata pine seedlings growing in a controlled growth cabinet was not significantly different over a range of constant PAR flux densities. Mean RUE’s were 3.22, 2.82 and 2.58 g MJ⁻¹ at 125, 250 and 500 µmol m⁻² s⁻¹ respectively.
In the second experiment, the novel use of a soil moisture modifier (ƒθ) to predict RUE of seedlings subjected to water stress proved successful within a limited range of soil water stress conditions. Measured seedling transpiration and stomatal conductance were closely correlated but seedling photosynthesis was less correlated with available soil water. This result suggests that photosynthesis was not coupled with stomatal conductance when PAR flux was 250 µmol m⁻² s⁻¹, which is well below saturating irradiance for C₃ plants.
Conclusions: The use of hybrid, quasi-mechanistic models to describe tree seedling growth has been seldom explored, which necessitated the development of novel experimental and analytical techniques for this study. These included a predictive model of germination decline at sub- and supra-optimal temperatures; a method for accurately estimating seedling dry weights under a range of PAR flux densities; and a novel method for estimating light interception by small seedlings.
The work reported in this thesis showed that existing hybrid models (the hydrothermal time germination model and the RUE model) can be adapted to model germination and growth of radiata pine seedlings under controlled environmental conditions. Nonetheless, further research is needed before the models can be confidently used as an alternative to conventional empirical models to model regeneration in “real-world” forests. Research priorities are the performance of hydrothermal germination models under variable field conditions, and the use of the soil moisture modifier for seedlings growing on a range of soil textures and under a range of PAR fluxes.||en