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Notes: The Arctic is often assumed to be an NH4+-dominated ecosystem. This review assesses the validity of this assumption. It also addresses the question of whether Arctic plant growth is limited by the ability to use the forms of nitrogen that are available. The review demonstrates that several sources of soil nitrogen are available to Arctic plants, including soluble organic nitrogen (e.g. glycine, aspartic acid and glutamic acid), NH4+ and NO−3. In mesic Arctic soils, soluble organic nitrogen is potentially more important than either NH+4 or NO−3. Many Arctic species are capable of taking up soluble organic nitrogen (either directly and/or in association with ectomycorrhizae), with the greatest potential for soluble organic nitrogen uptake being exhibited by deciduous species. The ability to take up soluble organic nitrogen may enable some Arctic plants to avoid nitrogen limitations imposed by the slow rate of organic matter decomposition. NO−3 is also present in many Arctic soils, especially calcareous soils and soils near flowing water, animal burrows and bird cliffs. Arctic species characteristic of mesic and xeric habitats are capable of taking up and assimilating NO−3. Even when present in lower concentrations in soils than NH+4, NO−3 is still an important source of nitrogen for some Arctic plants. Arctic-plants therefore have a variety of nitrogen sources available to them, and are capable of using those nitrogen sources. Taken together, these findings demonstrate that the Arctic is not an NH+4dominated ecosystem. Symbiotic fixation of atmospheric N2 does not appear to be an important source of nitrogen for Arctic plants. The reliance of Arctic plants on internal recycling of nitrogen substantially reduces their dependence on soil nitrogen uptake (this is particularly the case for slow-growing evergreens). Despite the high level of internal nitrogen recycling, Arctic plant growth remains limited by the low levels of available soil nitrogen. However, Arctic plant growth is not limited by an inability to utilize any of the available forms of nitrogen. The potential effects of climatic warming on nitrogen availability and use are discussed. The question of whether the Arctic ecosystem is uniquely different from temperate nitrogen-deficient ecosystems is also assessed.

Notes: In this study we assessed the inherent relative growth rate (RGR) under controlled environment conditions of 10 contrasting Acacia species from semi-arid and mesic environments. For several of the species, compound pinnate leaves produced early in the seedling stage, were gradually replaced by phyllodes (expanded petioles that form simple lamina). Other species either did not form phyllodes, or only did so to a minor degree by the end of the study. Phyllode production was dominant in the four slow-growing Acacia species from semi-arid environments (A. aneura, A. colei, A. coriacea and A. tetragonophylla), with leaf production being exclusive or dominant in five (A. dealbata, A. implexa, A. mearnsii, A. melanoxylon and A. irrorata) of the six faster-growing species from mesic environments. The exception was A. saligna which was fast growing but did produce phyllodes. From a carbon economy perspective, slow growth in the semi-arid species was not associated with lower net assimilation rates or less plant mass allocated to foliage. Rather, the primary factor associated with their slow growth was a smaller foliage area per unit foliage mass. This was true for comparisons based on the mean over all harvests or at set plant masses. The production of phyllodes by the semi-arid species substantially reduced foliage area per unit foliage mass, as this was lower for phyllodes than leaves in all species. To assess the impact that phyllode production had on ontogenetic changes in RGR, we modelled the situation where only leaves were formed. This analysis showed that changing from leaves to phyllodes substantially reduced the RGR. There was little difference in plant nitrogen concentration or the ratio of foliage nitrogen to plant nitrogen between the species. This resulted in foliage nitrogen productivity (dry mass gain per unit foliage nitrogen and time) being directly proportional to foliage area per unit foliage mass between species. We concluded that a smaller foliage area per unit foliage mass and phyllode production are the primary factors associated with lower RGR in contrasting Acacia species.

Notes: This study investigates the nitrogen economy of six altitudinally contrasting Poa species which differ in their relative growth rate (R). Two alpine (Poa fawcettiae and P. costiniana), one sub-alpine (P. alpina)and three temperate lowland species (P. pratensis, P. campressa and P. trivialis) were grown hydroponically under identical conditions in a growth room. The low R exhibited by the alpine species was associated with lower plant organic nitrogen concentration (np) and lower nitrogen productivity (Πp, amount of biomass accumulation per mol organic nitrogen and time). The differences in Πp between the alpine and lowland species did not appear to be due to differences in the carbon concentration or the proportion of total plant organic nitrogen allocated to the leaves, stems or roots. Variations in ΠP were also not due to variations in photosynthetic nitrogen use efficiency (ΨN, the rate of photosynthesis per unit organic leaf nitrogen) or shoot or root respiration rates per unit organic nitrogen (ΛSH and ΛR, respectively) per se. Rather, the lower Λp in the alpine species was probably due to a combination of small variations in several of the parameters (e.g. slightly lower ΨN, slightly higher ΛSH and ΛR, and slightly higher proportions of total plant organic nitrogen allocated to the roots). The alpine species exhibited lower organic acid and mineral concentrations. However, no differences in whole-plant construction costs (grams of glucose needed to synthesize one gram of biomass) were observed between She alpine and lowland Poa species. The lack of sub-stantial differences in ΨN between the alpine and lowland species contrasts with the large differences in ΨN between slow- and fast-growing lowland species that have been reported in the literature. The reasons for the unusually high ΨN values exhibited by the alpine Poa species are discussed.

Notes: The contribution of individual plant mitochondrial respiratory pathways to total respiration is commonly assessed by titration with specific inhibitors of different components in the branched electron transport chain. A pathway's contribution is equal to the activity when the other branch is blocked by an inhibitor multiplied by the degree (0-1.0) to which this activity is engaged when both pathways are operating. According to Bahr and Bonner (1973. J. Biol. Chem. 218: 3441–3445) the plot of the activities of identical titrations, one performed in the absence and the other in the presence of a specific inhibitor of the other branch of the respiratory chain, yields a straight line whose slope indicates the engagement of the titrated pathway during uninhibited respiration. An initial slope of zero may occur if electron flux is diverted between pathways during titrations. However, beyond the breakpoint (representing the point of pathway saturation), a straight line is obtained with a slope representing engagement. This technique assumes that the kinetics of inhibiting a specific component of the respiratory chain are independent of the absolute rate of electron flux through the total pathway. To test this assumption, the activity of respiratory pathways in isolated soybean (Glycine max [L]. Merr. cv. Stevens) mitochondria was titrated with specific inhibitors of the cytochrome and alternative oxidases. Under these conditions, the electron flux through a given pathway was manipulated by poising the rate of succinate oxidation with the succinate dehydrogenase inhibitor malonate. Construction of activity plots in the presence versus absence of malonate failed to result in straight lines for either KCN (when titrating the cytochrome pathway) or salicylhydroxamic acid (when titrating the alternative pathway). Rather, the resultant plots were always curvilinear whenever the activity in the presence of malonate divided by the activity in the absence of malonate was less than 1.0. In no case could the real engagement of the pathway be precisely estimated from the titration data. Titrations of cytochrome pathway activity in isolated potato tuber (Solanum tuberosum L. cv. Sabago and Canabex) mitochondria (which lack the alternative oxidase) showed that as the inhibitor concentration was increased, so did the reduction status of the ubiquinone pool, to a new steady state. The dependence of inhibition kinetics on the rate of flux through the pathway, and the increase in ubiquinone pool reduction upon KCN addition, are explained in terms of the elasticity of component enzymes as outlined in the theory of metabolic control analysis. The implications of this finding for the use of titrations to estimate engagement of plant respiratory pathways are discussed.