Hybrid physiological models are being increasingly used to assess productivity, carbon sequestration, water and nutrient use and environmental impacts of management decisions. Users include forest managers, politicians, environmental agencies and scientists. However a wider use of these models has been prevented as a result of an incomplete understanding of the mechanisms regulating carbon allocation, nutrient availability in soils and nutrient uptake by trees. On-going innovation in clonal forestry, genetic improvement and vegetation management techniques is also poorly represented in hybrid models. This thesis examines means to represent nutrition and genotype-nutrition interactions in productivity physiological hybrid models. Nutrient limitations and growth differences between genotypes were hypothesized to operate through key physiological processes: photosynthesis, carbon allocation and nutrient internal cycling. In order to accomplish the aims of the study both greenhouse and field experimentation were carried out. In a first experiment, responses of photosynthesis (A) to intercellular CO₂ concentration (Ci) were measured in a fast- and a slow-growing clone of Pinus radiata D. Don cultivated in a greenhouse in a factorial combination of nitrogen and phosphorus supply, and analyzed using the biochemical model of leaf photosynthesis described by Farquhar et al. (1980). There were significant positive linear relationships between the parameters, Vcmax, Jmax, Tp and both foliar nitrogen (Na) and phosphorus (Pa) concentration on an area basis. The study showed that the effects of nitrogen and phosphorus supply on photosynthesis were statistically independent and that the photosynthetic behaviour of the two clones was equivalent. In a similar study, gas exchange and chlorophyll fluorescence were simultaneously measured to determine internal transfer conductance (gm) based on the "constant J method". Transfer conductance may pose significant limitations to photosynthesis which may be differentially affected by nutrition and genotype in Pinus radiata. Values of gm were similar to those of stomatal conductance (gs) and their ratio (gm / gs) was not influenced by nutrient supply or clone being on average (±1 SE) 1.22 ±0.04. Relative mesophyll limitations (LM, 16%) to photosynthesis were marginally greater than those imposed by stomata (LS, 13%), and together smaller than the relative limitations posed to photosynthesis by biochemical processes (LB, 71%). The CO₂ concentration in the intercellular air spaces (Ci) was (±1 SE) 53 ±3 µmol mol-1 lower than in the atmosphere (Ca) while CO₂ concentration in the chloroplasts (Cc) was (±1 SE) 48 ±2 µmol mol-1 less than Ci. Values of LS, LM and LB and CO₂ diffusion gradients posed by gs (Ca-Ci) and gm (Ci-Cc) did not change with nutrient supply or clone. In a third experiment, one-year old Pinus radiata cuttings from four genotypes were cultivated in silica sand with a factorial combination of nitrogen (N0=1.43 and N1=7.14 mM) and phosphorus (P0=0.084 and P1=0.420 mM) supply for 24 months. N supply was enriched with ¹⁵N to 2.5⁰/₀₀ (labelled N) during the first year, then plants transferred to clean sand and cultivated for another year with ¹⁵N at levels close to natural abundance (0.3664899 atom percent ¹⁵N, δ¹⁵N 0.5115 ⁰/₀₀) provided by the source of N in nutrient solution applied during the second year. Recovery of labelled and unlabelled N was used to estimate N remobilization. N remobilization scaled with plant growth, N content and N and P supply. In relative terms, 65% of all stored N was remobilized in the high-nutrient supply regime compared to 42-48% at lower N and P addition rates. Most N remobilization occurred during spring-summer (77%), coincidently with the largest proportion of needle development (80%), indicating that N remobilization was driven by sink-strength. Foliage was by far the main source for internal cycling while roots were the main sink (40%). Clones exhibited differences in N remobilization capacity, but these differences were completely explained by the size of the N pool before remobilization took place, indicating that N remobilization performance was similar among clones. In a fourth study, four clones were cultivated in silica sand with a factorial combination of nitrogen and phosphorus supply for ten months, and patterns of carbon allocation examined using a carbon balance approach. Gross-primary productivity (GPP) scaled mainly with nitrogen but also with phosphorus supply. The fraction of GPP (GPP = ANPP + APR + TBCA) allocated to above-ground components (ANPP) increased with N and P supply at the expense of total-below ground C allocation (TBCA) with no apparent effect on the fraction of GPP partitioned to above-ground plant respiration (APR). Carbon use efficiency (NPP:GPP) scaled with nutrient supply, being 0.42 in the low-nutrient supply regime compared to 0.51 in the high-nutrient supply regime, suggesting that in poor fertility environments a larger proportion of the C budget is respired compared to the net productivity. Fast-growing clones allocated about 2-4% more carbon to above-ground components (ANPP) at the expense of carbon allocated below-ground (TBCA) with no effect on carbon respired above-ground (APR), indicating that faster-growing genotypes allocate more carbon to leaf area which may compound and increase overall GPP over time. The field component of this thesis was conducted in a subset of locations where ENSIS (formerly New Zealand Forest Research Institute) had established trials to test the influence of species, soil disturbance and plant nutrition on sustainability indicators. Plots were small in size (3 m × 3 m) with trees spaced at 0.5 m × 0.5 m (40 000 trees ha-1) with nine measurement trees surrounded by a two-row buffer. All sites were planted in winter 2001 and harvested in spring 2005. The aim of this pilot study was to examine patterns of carbon allocation during the fourth year after planting in control and fertilized mini-plots of Pinus radiata in five sites with contrasting climate and soil conditions in the South Island of New Zealand. The study showed that the fraction of gross-primary productivity allocated belowground increased as the soil C:N ratio increased. However, these results should be interpreted with caution due to the unusual nature of the trial and the reduced number of sites studied. Two existing physiological models were selected for the discussion in this thesis (3-PG, Landsberg and Waring 1997; canopy net carbon exchange model, Whitehead et al. 2002). Potential improvements for the nutritional component of 3-PG comprise: accounting for reductions in carbon use efficiency (NPP:GPP) in poor-fertility environments, adding a preliminary fertility modifier (FN, 0-1) driven by soil C : N ratio and soil N, adding a preliminary relationship between carbon allocation to roots and the soil C : N ratio and representing faster-growing genotypes by increasing their leaf area but not their photosynthetic performance. The canopy net carbon exchange model (NCE) combines the coupled model of leaf photosynthesis - stomatal conductance described by Leuning (1995) with canopy structure and a water balance model to scale carbon assimilation from leaves to canopies. Potential improvements to account for nutrient deficiencies in the leaf model by Leuning (1995), comprise using nutrient ratios to discriminate nitrogen (Na/Pa < 23 mol mol-1) from phosphorus deficiencies (Na/Pa > 23 mol mol-1), adding relationships between photosynthetic model parameters Vcmax and Jmax to Pa, and correcting the estimation of photosynthetic parameters Vcmax and Jmax by accounting for transfer conductance (gm). The canopy net carbon exchange model may be also modified to account for carbon-use efficiency, carbon allocation to roots and genotype in a similar form to that proposed for 3-PG. The results previously outlined provide a preliminary framework to represent tree and soil nutrition in physiological hybrid productivity models.

No description available.

More than 20 million clonal loblolly pines have been planted throughout the southeastern United States. Fertilizer has been applied to more than 6.5 million hectares of plantations to alleviate deficiencies of nitrogen and phosphorus that limit growth. Because cloning loblolly pine in large numbers has only become possible in the last decade, it is unknown how clones may respond differently to fertilizer application. Growth, growth efficiency, and biomass partitioning responses to fertilizer application were investigated among 25 clones planted in the Virginia Piedmont. Closely related clones varied in their fertilizer stem volume responses, but not enough to be statistically significant (p = 0.11). Clones varied in growth efficiency and partitioning to individual tissues, but clone-by-fertilizer interactions were not observed. Clonal variability was observed in root morphology, and maximum rooting depth showed a significant clone-by-fertilizer interaction. Clones with rapid growth rates can be selected with a range of other desirable traits. Short-term (i.e. weeks) responses to fertilization are often inconsistent with long-term (i.e. years) responses, but are critical to understanding growth responses. We investigated carbon allocation in two full-sibling clones of loblolly pine under two levels of fertilizer application over four months in a greenhouse. Using monthly harvests of some trees and ecophysiological measurements throughout, we determined carbon allocation on a monthly scale. In response to fertilizer application, both clones reduced allocation belowground and increased allocation to foliage to some extent, increasing whole-canopy photosynthetic capacity. However, these changes in allocation were ephemeral. By the end of the experiment, root-shoot ratios were no longer significantly affected by fertilizer application. Clones had allocation patterns distinct from one another, with one allocating more belowground and the other allocating more to stem mass. While their overall growth responses to fertilizer application were similar, the physiological mechanisms that resulted in these responses were different between clones. Results of the two studies indicate that while fertilizer responses may not need to be included when testing clones for deployment, knowledge of the fertilizer responses of widely-deployed clones would offer forest managers opportunities to apply clone-specific precision-silvicultural systems to optimize growth rates and manage for a range of products. / Ph. D.

Soils contain the largest terrestrial pool of carbon (C), but the magnitude and distribution of the soil C sink may be sensitive to climate change. My dissertation aims to identify key processes that mediate patterns of belowground carbon storage across the globe and quantify the effect of environmental perturbations associated with global change on existing soil carbon stocks in peatland ecosystems. Using meta-analysis, I show that the relationship between plant growth, C allocation, and soil nutrient availability varies on a global scale and high-latitude ecosystems allocate >60% of fixed C to belowground structures. As high latitude ecosystems are warming faster than the global mean, the future of this belowground C store is potentially sensitive to climate change. In high latitude ecosystems in particular, I further show that belowground warming increases the rate of peatland carbon dioxide (CO2) and methane (CH4) losses, although CH4 emissions are more sensitive to warming than CO2 emissions, which is likely to shift the nature of greenhouse gas emissions and increase the importance of CH4 as a radiative forcing agent in the near-term. I also use a natural peatland water table gradient to identify the effect of water table reduction on peatland C and N cycling and find that microbial community shifts in C and N demand may attenuate production of C-degrading enzymes and C mineralization in the presence of plant roots and in areas with low water tables. Together, my dissertation work highlights the important role of belowground plant and microbial processes in high latitude ecosystems, and identifies the potential influence of factors associated with global change on belowground C and nutrient cycling.

Patterns of woody-plant mortality have been linked to global-scale environmental changes, such as extreme drought, heat stress, more frequent and intense fires, and episodic outbreaks of insects and pathogens. Although many studies have focussed on survival and mortality in response to specific physiological stresses, little attention has been paid to the role of genetic heritability of traits and local adaptation in influencing patterns of plant mortality, especially in non-native species. Tamarix spp. is a dominant, non-native riparian tree in western North America that is experiencing dieback in some areas of its range due to episodic herbivory by the recently introduced northern tamarisk leaf beetle (Diorhabda carinulata). We propose that genotype x environment interactions largely underpin current and future patterns of Tamarix mortality. We anticipate that (i) despite its recent introduction, and the potential for significant gene flow, Tamarix in western North America is generally adapted to local environmental conditions across its current range in part due to hybridization of two species; (ii) local adaptation to specific climate, soil and resource availability will yield predictable responses to episodic herbivory; and (iii) the ability to cope with a combination of episodic herbivory and increased aridity associated with climate change will be largely based on functional tradeoffs in resource allocation. This review focusses on the potential heritability of plant carbon allocation patterns in Tamarix, focussing on the relative contribution of acquired carbon to non-structural carbohydrate (NSC) pools versus other sinks as the basis for surviving episodic disturbance. Where high aridity and/or poor edaphic position lead to chronic stress, NSC pools may fall below a minimum threshold because of an imbalance between the supply of carbon and its demand by various sinks. Identifying patterns of local adaptation of traits related to resource allocation will improve forecasting of Tamarix population susceptibility to episodic herbivory.

6 Abstract This study aimed to compare the mycorrhizal fungal communities inhabiting the roots of invasive Pinus strobus L. and native Pinus sylvestris L. We also compared carbon allocation into ectomycorrhizal fungal (EcMf) and other structures of the two pine species. The aim was to assess the influence of mycorrhiza on the invasive potential of P. strobus in the protected areas of National Park Bohemian Switzerland. The two field experiments were conducted on three locations of each species. We estimated the EcM extramatrical mycelium (EMM) production by measuring the ergosterol content in sterile sand filled mesh-bags. Next measured variables were: biomass of ectomycorrhizal and saprotrophic sporocarps, fine roots biomass and leaf litter biomass to compare the one season production of each measured variables. The results revealed a major difference in EcM sporocarps production, whereas on the P. sylvestris sites was the production 100% higher. Same results came from the fine roots measurements: P. sylvestris had a higher fine roots production, which may be also related with the production of EMM, which was about 60% higher as well. The EcMf species richness on the P. strobus root-tips was as high as the native pine, but the species composition was different. The P. strobus prefers EcMf species with...

FTIR spectroscopy as molecular fingerprint has been used to assess macromolecular and ele-mental stoichiometry as well as growth rates of phytoplankton cells. Chemometric models have been developed to extract quantitative information from FTIR spectra to reveal macro-molecular composition (of proteins, carbohydrates and lipids), C:N ratio, and growth potential. In this study, we tested these chemometric models based on lab-cultured algal species in mon-itoring changes of phytoplankton community structure in a hypertrophic lake (Lake Auensee, Leipzig, Germany), where a seasonal succession of spring green algal bloom followed by cya-nobacterial dominance in summer can be commonly observed. Our results demonstrated that green algae reacted to environmental changes such as nitrogen limitation (due to imbalanced nitrogen and phosphorus supply) with restricted growth by changing carbon allocation from protein synthesis to storage carbohydrates and/or lipids, and increased C:N ratio. By contrast, cyanobacteria proliferated under nitrogen limiting conditions. Furthermore, the FTIR-based growth potential of green alga matched well with the population biomass determined by the Chl-a concentration. However, the predicted growth potential based on FTIR spectroscopy cannot describe the realistic growth development of cyanobacteria in this lake. These results revealed that green algae and cyanobacteria have different strategies of C-allocation stoichi-ometry and growth patterns in response to environmental changes. These taxon-specific re-sponses may explain at a molecular level why green algae bloomed in the spring under condi-tions with sufficient nutrient, lower pH and lower water temperature; while cyanobacteria overgrew green algae and dominated in the summer under conditions with limited nutrient availability, higher pH and higher water temperature. In addition, the applicability of these chemometric models for predicting field cyanobacterial growth is of limited value. This may be attributed to other special adaptation properties of cyanobacterial species under stress growth conditions. We used flow cytometry to isolate functional algal groups from the water samples. Despite some drawbacks of the flow cytometry combined FTIR spectroscopy tech-nique, this method provides prospects of monitoring water quality and early warning of harmful algal blooms.

Die Rotbuche (Fagus sylvatica L.) ist die bestimmende Baumart der potentiell natürlichen Vegetation in den Wäldern Mittel- und Westeuropas die ökonomisch bedeutsamste Laubbaumart Deutschlands. Obwohl diese spät-sukzessionelle Baumart über eine hohe physiologische Toleranz gegenüber einem weiten Spektrum klimatischer Wuchsbedingungen verfügt, wird die Buche gegenüber anderen temperaten Laubbaumarten als relativ trockensensitiv eingeschätzt. Da im Zuge des globalen Klimawandels mit einer Verschlechterung der klimatischen Wasserbilanz und mit einer Zunahme sommerlicher Trockenperioden gerechnet wird, wird die zukünftige Rolle der Rotbuche in der europäischen Forstwirtschaft derzeit intensiv diskutiert. Diese Studie hatte zum Ziel, hydrologische und klimatische Einflüsse auf die Produktivität und die Vitalität der Rotbuche zu untersuchen. Hierdurch sollen grundlegende Mechanismen der Trockenstressantwort bei dieser trocken-sensitiven Art identifiziert, und Rückschlüsse auf zukünftige Klimaantworten von Buchenbeständen ermöglicht werden. Zu diesem Zweck wurde die ober- und unterirdische Biomasseproduktion von 12 Buchenaltbeständen im Norddeutschen Tiefland entlang eines natürlichen Niederschlagsgradienten (543-816 mm a-1) auf einheitlichem geologischen Substrat ermittelt. Um den zusätzlichen Einfluss der Wasserspeicherkapazität der Böden zu berücksichtigen, wurden Paare von Buchenbeständen untersucht, die unter nahezu identischen klimatischen Bedingungen, jedoch auf Böden unterschiedlicher Textur (sandige versus lehmig-sandige Böden) stockten. Einflüsse der Wasserverfügbarkeit und klimatischer Variationen auf das Wachstum wurden untersucht unter Berücksichtigung (i) der gesamten ober- und unterirdischen Biomasseproduktion, (ii) der Dynamik von Ressourcen-Allokation und Kohlenstoff-Partitionierung, sowie (iii) der Morphologie wasseraufnehmender und -abgebender Oberflächen. Unerwarteterweise zeigte sich die gesamte Produktivität von Buchen-Altbeständen nur geringfügig von Veränderungen der hydrologischen Regime entlang des Gradienten beeinflusst. Trotz deutlicher Unterschiede in der jährlichen Wasserverfügbarkeit nahmen die oberirdische und die gesamte Biomasseproduktion auf den trockeneren Flächen des Transektes nicht ab. Allerdings führten ausgeprägte früh-sommerliche Wasserdefizite (in den Monaten Juni und Juli) zu deutlichen Einbußen der oberirdischen Biomasseproduktion, und insbesondere der Stammholzproduktion. Entlang des untersuchten Gradienten konnte eine ausgeprägte, kontinuierliche Verschiebung der Allokationsmuster festgestellt werden: Mit abnehmender Wasserverfügbarkeit nahm die Feinwurzelproduktion zu und das Verhältnis von oberirdischer:unterirdischer Biomasseproduktion ab. Anders als oberirdische Komponenten zeigte die Feinwurzelproduktion eine hohe Sensibilität gegenüber Unterschieden hinsichtlich hydrologischer Regime. In Übereinstimmung mit der Optimalitätstheorie der pflanzlichen Ressourcennutzung konnte dieses Allokationsverhalten in sowohl in Reaktion auf veränderte Niederschläge, als auch in Antwort auf auch veränderte Wasserspeicherkapazitäten beobachtet werden. Allokative Anpassungsmechanismen an Wassermangel wurden im Feinwurzelbereich zusätzlich durch morphologische Plastizität (Zunahme im Verhältnis von Oberfläche: Biomasse) und durch Regulierung der räumlichen Verteilung (zunehmende Konzentrierung von Feinwurzeln in der organischen Auflage) komplementiert. Im Gegensatz zu diesen komplexen unterirdischen Trockenheits-Antworten konnten keinerlei Anpassungen der Blattmorphologie an veränderte hydrologische Bedingungen festgestellt werden. Neben Reaktionen auf Wasserverfügbarkeit wurde die Fruchtbildung als zweiter wesentlicher Einfluss auf das Allokationsverhalten der Buche erkannt. Eine deutliche Ressourcen-Allokation zu Gunsten der Fruchtentwicklung beeinträchtigte maßgeblich das oberirdische vegetative Wachstum, insbesondere den Stammholzzuwachs. Auf Grund einer hohen Attraktionsstärke der Früchte gegenüber C und N führte zunehmende Fruktifizierung auch zu einer Gewichts- (und Größen-) Abnahme der Einzelblätter und somit zu reduzierter Bildung von Blattmasse und Bestandesblattfläche (LAI). Neben dieser Abnahme an assimilierender Blattoberfläche führte auch eine deutliche Senkung der Blatt-Stickstoffgehalte in Folge der reproduktiven Ressourcenwidmung mutmaßlich zu einer Verschlechterung der C-Bilanz, sowohl im Mast- als auch im Folgejahr. Eine Analyse klimatischer Einflussfaktoren auf das Mastverhalten legt nahe, dass die Blütenbildung der Buche durch Überschreitung eines Schwellenwertes der Kohlenstoffassimilation im Frühsommer (Juni-Juli) induziert wird. Sofern diese Schlüsse zutreffen, unterliegt das zeitliche Muster der Fruktifikations-Antwort auf Witterungsauslöser einer Rückkopplungskontrolle durch pflanzliche Stickstoff-Dynamik. Vor dem Hintergrund anhaltend erhöhter Stickstoffdepositionen ergäbe sich aus diesem Mechanismus eine zusätzliche Belastung für das zukünftige vegetative Wachstum der Buche. Es ist anzunehmen, dass die in dieser Studie belegte hohe allokative Plastizität in Altbäumen Fagus sylvatica dazu befähigt, ihre hohe Konkurrenzkraft in einem breiten Spektrum hydrologischer Regime zu entfalten. Darüber hinaus werden die hier dargestellten Mechanismen einer langfristigen Trockenheitsanpassung mutmaßlich zu einer gesteigerten Resistenz und Resilienz von Buchen-Altbeständen gegenüber Ereignissen extremer Sommertrockenheit beitragen.

Fotossíntese é o processo biofísico pelo qual energia luminosa é transformada em energia química armazenada em compostos de carbono. A taxa fotossintética instantânea possui um forte padrão assintótico em resposta ao incremento da intensidade luminosa, porém quando integramos a fotossíntese em escalas espaciais e temporais maiores, observa-se um padrão linear de resposta entre radiação interceptada e produção. Esta abordagem permitiu o surgimento de modelos baseados nas taxas de conversão de energia radiante em biomassa seca, ou eficiência do uso da luz (&epsilon;). Valores publicados para o Eucalyptus estão na faixa de 0,5-2,5 g MJ-1, porém se faz necessário um entendimento mais profundo a respeito da sensibilidade destes valores às flutuações do clima e sua sazonalidade. Para isso, as taxas de crescimento, uso e eficiência do uso da luz foram monitoradas quinzenalmente durante 16 meses em parcelas de 18 clones de Eucalyptus, dos 1,3 aos 2,7 anos de idade. Foram testadas as hipóteses de que a produção de madeira aumentaria em função de incrementos no uso e/ou eficiência de uso da luz, assim como estes valores aumentariam respectivamente com incrementos no índice de área foliar e por uma alocação de carbono para o fuste, respectivamente. Os clones apresentaram uma grande amplitude de produtividade (9,9-22,7 Mg ha-1 ano-1) e arquiteturas de copa, capturando entre 65-95% da radiação incidente. Tais valores resultaram em uma eficiência do uso da luz média de 1,5 g MJ-1, variando entre 0,16-3,14 g MJ-1. Apesar de patamares distintos, os valores de eficiência de uso dos clones oscilaram de maneira similar, de modo que a radiação incidente foi a principal variável afetando a eficiência de uso da luz, estando &epsilon; positivamente relacionada a variáveis que expressam períodos de maior disponibilidade hídrica e negativamente relacionado a períodos de menor disponibilidade. Maiores valores de índice de área foliar efetivo (Le) acarretaram em maior interceptação de luz, porém as distintas arquiteturas de copa revelaram diferentes estratégias de captura de luz (0,3 < &kappa; < 0,6). Apesar de uma maior interceptação, não houve correlação significativa com a produtividade, no entanto observou-se uma forte correlação entre eficiência do uso da luz e crescimento em madeira, resultado de uma maior alocação para o fuste. Apesar de evidenciar a relação entre alocação e eficiência, existem outros mecanismos associados às alterações observadas em &epsilon; que apenas uma caracterização completa dos fluxos de carbono pode elucidar. / Photosynthesis is the biophysical process by which light energy is converted into chemical energy stored in carbon compounds. The instantaneous photosynthetic rate has a strong asymptotic pattern in response to increases in light intensity, however when we integrate photosynthesis in larger spatial and temporal scales, there is a linear pattern of response between intercepted radiation and production. This approach has allowed the appearance of models based on radiant energy conversion rates into dry biomass, or light use efficiency (&epsilon;). Published values for Eucalyptus range from 0.5 to 2.5 g MJ-1, but a deeper understanding of the sensitivity of these values to climate fluctuations and seasonality is necessary. For this reason, wood growth rates, light use and efficiency were monitored every two weeks for 16 months at 18 Eucalyptus clones plots, from 1.3 to 2.7 years of age. Our hypothesis was that wood production would be positively related to light use and efficiency, as well these values would increase respectively with increases in leaf area index and carbon allocation to the stem. Clones showed a wide range of productivity (9.9 to 22.7 Mg ha-1 yr-1) and canopy architectures, capturing between 65-95% of incident radiation. Such values resulted in an average light use efficiency of 1.8 g MJ-1, ranging from 0.16 to 3.14 g MJ-1. Although different levels, light use efficiency values for the clones fluctuated similarly. Incident radiation was the main variable affecting the efficiency of dry matter conversion, and &epsilon; values were positively related variables expressing periods of greater water availability and negatively related to periods of lower availability. Larger effective leaf area index (Le) values resulted in higher light interception, but the different canopy architectures revealed different light capture strategies (0.3 < &kappa; < 0.6). Despite a higher interception, there was no significant correlation with productivity; however there was a strong correlation between light use efficiency and wood growth, as a result of increased allocation to the stem. While evidencing the relationship between allocation and efficiency, there are other mechanisms associated with changes in &epsilon; observed that only one full characterization of the carbon fluxes can elucidate.

Mixed-species plantations of eucalypts and acacias have the potential to improve stand productivity over that of respective monocultures through the facilitative effect of nitrogen-fixation by acacias, and increased resource capture through above- and belowground stratification. However, growth in mixed-species plantations may not be improved compared to that of monocultures when competitive interactions outweigh the effects of improved nutrient availability and resource capture. Careful selection of sites and species is therefore critical to successfully improving stand productivity using mixed-species plantations. This study set out to examine some of the processes and interactions that occur in mixed-species plantations, and the effect nutrient and water availability can have on the growth of mixtures. In three out of four mixed-species field trials examined in this study, growth was not increased in mixtures compared to monocultures. However, in the fourth field trial, heights, diameters, stand volume and aboveground biomass were higher in mixtures of E. globulus and A. mearnsii from 3-4 years after planting. The range in outcomes from mixing species in these four trials shows that a fundamental understanding of the underlying processes is required to enable a greater predictive capacity for the circumstances under which mixtures will be successful. Therefore the growth dynamics, processes and interactions were examined in the mixtures of E. globulus and A. mearnsii. The difference in productivity between mixtures and monocultures in this trial increased with time up to age 11 years, when 1:1 mixtures contained twice the aboveground biomass of E. globulus monocultures. The positive growth response of trees in mixture compared to monocultures was the result of accelerated rates of nutrient cycling, a shift in C allocation and reductions in light competition through canopy stratification. Nitrogen contents of foliage and soil clearly showed that A. mearnsii influenced the N dynamics in this trial. If these changes in N contents were due to N fixation by A. mearnsii, then about 51 and 86 kg N ha-1 yr-1 was fixed in the 1:1 mixtures and A. mearnsii monocultures, respectively. Nitrogen fixation was also examined using the natural abundance method. The delta15N values of foliage collected at 10 years were grouped according to the mycorrhizal status of the host plant. Therefore the discrimination of 15N during transfer from mycorrhizae to the host plant appeared to vary with mycorrhizal status, and the natural abundance of 15N was not used to quantify N fixation. Rates of N and P cycling in litterfall were significantly higher in stands containing at least 25% A. mearnsii (more than 31 kg N ha-1 yr-1 and more than 0.68 kg P ha-1 yr-1) compared to E. globulus monocultures (24 kg N ha-1 yr-1 and 0.45 kg P ha-1 yr-1). Rates of litter decomposition and N and P release were about twice as high in 1:1 mixtures compared to E. globulus monocultures and were even higher in A. mearnsii monocultures. It is therefore important to select N-fixing species that are capable of cycling nutrients quickly between the plant and soil, and that have readily decomposable litter. The total belowground C allocation was not significantly different between mixtures and monocultures (14 to 16 Mg C ha-1 yr-1). However, since aboveground net primary production was greater in 1:1 mixtures, the changes in nutrient availability appears to have increased total productivity (both above- and belowground), and reduced the proportion of C allocated belowground in mixtures compared to E. globulus monocultures. In a pot trial containing mixtures of E. globulus and A. mearnsii both species grew larger in mixture than in monoculture at low N levels, and mixtures were more productive than monocultures. However, at high N levels, E. globulus suppressed A. mearnsii and mixtures were less productive than E. globulus monocultures. Similar effects were found for high and low levels of P. Therefore resource availability can have a strong influence on the interactions and growth of mixtures. The productivity of mixtures may only be increased on sites where the resource for which competition is reduced in mixture is a major limiting growth resource. For example, if N is not a limiting growth factor then an increase in N availability from N-fixation may not increase growth, and the N-fixing species may compete for other resources such as soil P, moisture or light. This study has shown that mixtures containing a N-fixing trees and a non-N-fixing trees can be more productive than monocultures, but that this increase in productivity will only occur on certain sites. Examination of the growth, interactions and processes that occurred in mixtures in this study provide useful information that can aid the selection of species combinations and sites.