Over large areas of Europe, coniferous monocultures are being transformed into mixed forests by the re-introduction of broadleaf tree species belonging to the potential natural vegetation. One important species of interest in this changing forest policy is European beech (Fagus sylvatica). However, at present, this forest management directive has ignored potential adverse effects of global climate change on wide-spread re-introduction of beech to these areas. Average global surface temperatures have risen by ap-

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The heat wave of summer 2003 was the largest and the most persistent ever experienced in Central Europe and has fuelled concern about the effects of climate change on European ecosystems. Since forests constitute the most important European ecosystems, in this review article we assess current knowledge on the effects of heat and drought on key metabolic processes for growth and productivity of forest trees. In particular, the general consequences of heat and drought on (1) photosynthesis and respiration at the cellular and community level, and (2) on nutrient uptake, partitioning and competition for nutrients are summarized. The latter are a major sink for photosynthetic energy and, therefore, are indirectly but strongly connected to the performance of photosynthesis. In addition, the interaction of heat and drought with stress compensation mechanisms and emission of biogenic volatile organic compounds (BVOC) are discussed, since these processes are directly connected to carbon metabolism. Effects on the emission of BVOC are also included because they constitute an important feedback mechanism on ozone formation and, thus, on atmospheric pollution. As far as available, data collected during the 2003 heat wave are included and discussed.

The molecular and physiological responses of gray poplar (Populus 3 canescens) following root hypoxia were studied in roots and leaves using transcript and metabolite profiling. The results indicate that there were changes in metabolite levels in both organs, but changes in transcript abundance were restricted to the roots. In roots, starch and sucrose degradation were altered under hypoxia, and concurrently, the availability of carbohydrates was enhanced, concomitant with depletion of sucrose from leaves and elevation of sucrose in the phloem. Consistent with the above, glycolytic flux and ethanolic fermentation were stimulated in roots but not in leaves. Various messenger RNAs encoding components of biosynthetic pathways such as secondary cell wall formation (i.e. cellulose and lignin biosynthesis) and other energy-demanding processes such as transport of nutrients were significantly down-regulated in roots but not in leaves. The reduction of biosynthesis was unexpected, as shoot growth was not affected by root hypoxia, suggesting that the up-regulation of glycolysis yields sufficient energy to maintain growth. Besides carbon metabolism, nitrogen metabolism was severely affected in roots, as seen from numerous changes in the transcriptome and the metabolome related to nitrogen uptake, nitrogen assimilation, and amino acid metabolism. The coordinated physiological and molecular responses in leaves and roots, coupled with the transport of metabolites, reveal important stress adaptations to ensure survival during long periods of root hypoxia.Higher plants are aerobic organisms and depend on the availability of O 2 . A lack of O 2 in the rhizosphere affects the maintenance of numerous pathways and is therefore an important environmental stress for vascular plants (Drew, 1997). Plant adaptations to O 2 deprivation include avoidance strategies at the morphological level and physiological tolerance mechanisms (Bailey-Serres and Voesenek, 2008). One of the major cellular pathways dependent on O 2 is mitochondrial respiration. In order to maintain energy generation under conditions of decreased O 2 availability, plants switch from respiration to fermentative metabolism. Fermentation allows regeneration of NAD + in the absence of respiration, thereby maintaining glycolysis and the generation of ATP under anaerobic conditions. As an initial reaction to O 2 deprivation, many plants activate lactic acid fermentation. As generation of lactic acid causes a decrease in cytosolic pH (Roberts et al., 1984), which reduces the activity of the responsible enzyme, lactate dehydrogenase (LDH; Hanson and Jacobson, 1984), lactic acid fermentation is followed by alcoholic fermentation (Davies et al., 1974;Roberts et al., 1984). The significantly lower energy yield of alcoholic fermentation, compared with mitochondrial respiration, causes an energy crisis in anaerobic tissues (Bailey-Serres and Voesenek, 2008).A high rate of fermentation increases the demand for carbohydrates, leading to the hypothesis that carbohydrate supply becomes c...

During the vegetation periods 1994 and 1995, net uptake of nitrate and ammonium by roots of adult spruce (Picea abies (L.) Karst) and beech (Fagus sylvatica L.) trees was studied at a field site exposed to high loads of N (' Ho$ glwald ', Germany). In addition, uptake experiments were carried out under controlled conditions with young spruce and beech trees grown at normal N supply.In the field, nitrate was not taken up by the roots of spruce trees in appreciable amounts. This was also true for beech except during September 1995. Apparently, beech trees was capable of taking up nitrate, but the environmental condition prevailing at the field site usually prevented net uptake. Net uptake of ammonium in both tree species showed a seasonal course, with maximum rates in mid summer. Rates of ammonium uptake by both species correlated with soil temperature at the field site.Laboratory experiments on the influence of root temperature on uptake of nitrate indicated that uptake rates at temperatures found in the field were low compared with the uptake capacity at optimum temperature. At temperatures of 10 and 15 mC, frequently found in the soil at the field site, net uptake of nitrate by spruce and beech amounted to c. 16 % and 11 %, respectively, of maximum uptake at 25 mC. By contrast, net uptake of ammonium at 10 mC reached 73 % and 31 % of the maximum uptake for spruce and beech trees, respectively. Independent of temperature, rates of nitrate uptake were considerably lower than those of ammonium. In young spruce and beech trees, net uptake of nitrate was significantly inhibited by ammonium at nitrate : ammonium ratios found in the soil solution at the forest site. Preincubation of roots of both species, with amino acids present in the phloem of adult trees at the field site, led to an increase in the amino acid pool in the roots. For spruce trees a correlation between inhibition of uptake of nitrate and enrichment of the roots with the amino compounds Glu, γ-amino butyric acid (Gaba), Gln, and Asn was observed. In beech trees, enrichment of Asp and Gln in the roots correlated with a decrease in net uptake of nitrate. The results of laboratory experiments on the effects of temperature, the nitrate to ammonium ratio in the nutrient solution, and amino acid enrichment in the roots are discussed with special emphasis on the patterns of net uptake of ammonium and nitrate observed in the field.Key words : Nitrate uptake, ammonium uptake, soil temperature, nitrate to ammonium ratio, amino acids. In forest ecosystems, nitrate and ammonium are the most abundant N compounds available to the roots * To whom correspondence should be addressed. E-mail : here!sun2.ruf.uni-freiburg.de of trees. Total amounts of nitrate and ammonium, and the ratio of nitrate to ammonia depend on quality and quantity of N input and on the balance of ammonification, nitrification, immobilization and denitrification processes in the soil (Haynes & Goh,