We evaluated the carbon budget of coarse woody debris (CWD) in a temperate broad‐leaved secondary forest. On the basis of a field survey conducted in 2003, the mass of CWD was estimated at 9.30 tC ha−1, with snags amounting to 60% of the total mass. Mean annual CWD input mass was estimated to be 0.61 tC ha−1 yr−1 by monitoring tree mortality in the forest from 1999 to 2004. We evaluated the CWD decomposition rate as the CO2 evolution rate from CWD by measuring CO2 emissions from 91 CWD samples (RCWD) with a closed dynamic chamber and infrared gas analysis system. The relationships between RCWD and temperature in the chamber, water content of the CWD, and other CWD characteristics were determined. By scaling the measured RCWD to the ecosystem, we estimated that the annual RCWD in the forest in 2003 was 0.50 tC ha−1 yr−1 or 10%–16% of the total heterotrophic respiration. Therefore, 0.11 tC ha−1 yr−1 or 7% of the forest net ecosystem production was sequestered by CWD. In a young forest, in which CWD input and decomposition are not balanced, the CWD carbon budget needs to be quantified for accurate evaluation of the forest carbon cycle and NEP.

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A B S T R A C TTo separate CO 2 efflux from roots (R r ) and soil (R s ), we developed a system to measure R r continuously. Using this system, seasonal variation in R r was obtained in a temperate forest in Japan. We measured R s , CO 2 efflux from mineral soil (R m ) and environmental factors simultaneously, and the characteristic and seasonality of R r were analysed in comparison with R s . R r and R s showed different responses to soil water content: R s decreased with decreasing soil water content, whereas R r peaked at relatively low soil water content. R r /R s decreased from 64.8% to 27.3% as soil water content increased from 0.075 to 0.225 cm cm −3 . The relationship between respiration and temperature appears to change seasonally in response to phenological and biological factors. R r showed clear seasonal variation as a function of soil temperature. During the growing period, R r exhibited a higher rate at the same soil temperature than during other periods, which may be due to phenological influences such as fine root dynamics. R s decreased during the summer despite high soil temperatures. The seasonal peak for R s occurred earlier than that for soil temperature. R r /R s ranged between 25% and 60% over the course of the year.

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Main conclusion Eucalyptus camaldulensis EcDQD/SDH2 and 3 combine gallate formation, dehydroquinate dehydratase, and shikimate dehydrogenase activities. They are candidates for providing the essential gallate for the biosynthesis of the aluminum-detoxifying metabolite oenothein B. Abstract The tree species Eucalyptus camaldulensis shows exceptionally high tolerance against aluminum, a widespread toxic metal in acidic soils. In the roots of E. camaldulensis, aluminum is detoxified via the complexation with oenothein B, a hydrolyzable tannin. In our approach to elucidate the biosynthesis of oenothein B, we here report on the identification of E. camaldulensis enzymes that catalyze the formation of gallate, which is the phenolic constituent of hydrolyzable tannins. By systematical screening of E. camaldulensis dehydroquinate dehydratase/shikimate dehydrogenases (EcDQD/SDHs), we found two enzymes, EcDQD/SDH2 and 3, catalyzing the NADP+-dependent oxidation of 3-dehydroshikimate to produce gallate. Based on extensive in vitro assays using recombinant EcDQD/SDH2 and 3 enzymes, we present for the first time a detailed characterization of the enzymatic gallate formation activity, including the cofactor preferences, pH optima, and kinetic constants. Sequence analyses and structure modeling suggest the gallate formation activity of EcDQD/SDHs is based on the reorientation of 3-dehydroshikimate in the catalytic center, which facilitates the proton abstraction from the C5 position. Additionally, EcDQD/SDH2 and 3 maintain DQD and SDH activities, resulting in a 3-dehydroshikimate supply for gallate formation. In E. camaldulensis, EcDQD/SDH2 and 3 are co-expressed with UGT84A25a/b and UGT84A26a/b involved in hydrolyzable tannin biosynthesis. We further identified EcDQD/SDH1 as a “classical” bifunctional plant shikimate pathway enzyme and EcDQD/SDH4a/b as functional quinate dehydrogenases of the NAD+/NADH-dependent clade. Our data indicate that in E. camaldulensis the enzymes EcDQD/SDH2 and 3 provide the essential gallate for the biosynthesis of the aluminum-detoxifying metabolite oenothein B.