No abstract

There is a need to reappraise the effects of UV-B radiation on plant morphology in light of improved mechanistic understanding of UV-B effects, particularly elucidation of the UV RESISTANCE LOCUS 8 (UVR8) photoreceptor. We review responses at cell and organismal levels, and explore their underlying regulatory mechanisms, function in UV protection and consequences for plant fitness. UV-induced morphological changes include thicker leaves, shorter petioles, shorter stems, increased axillary branching and altered root:shoot ratios. At the cellular level, UV-B morphogenesis comprises changes in cell division, elongation and/or differentiation. However, notwithstanding substantial new knowledge of molecular, cellular and organismal UV-B responses, there remains a clear gap in our understanding of the interactions between these organizational levels, and how they control plant architecture. Furthermore, despite a broad consensus that UV-B induces relatively compact architecture, we note substantial diversity in reported phenotypes. This may relate to UV-induced morphological changes being underpinned by different mechanisms at high and low UV-B doses. It remains unproven whether UV-induced morphological changes have a protective function involving shading and decreased leaf penetration of UV-B, counterbalancing trade-offs such as decreased photosynthetic light capture and plant-competitive abilities. Future research will need to disentangle seemingly contradictory interactions occurring at the threshold UV dose where regulation and stress-induced morphogenesis overlap.

Cloud cover increases the proportion of diffuse radiation reaching the Earth's surface and affects many microclimatic factors such as temperature, vapour pressure deficit and precipitation. We compared the relative efficiencies of canopy photosynthesis to diffuse and direct photosynthetic photon flux density (PPFD) for a Norway spruce forest (25-year-old, leaf area index 11 m 2 m À2 ) during two successive 7-day periods in August. The comparison was based on the response of net ecosystem exchange (NEE) of CO 2 to PPFD. NEE and stomatal conductance at the canopy level (G canopy ) was estimated from halfhourly eddy-covariance measurements of CO 2 and H 2 O fluxes. In addition, daily courses of CO 2 assimilation rate (A N ) and stomatal conductance (G s ) at shoot level were measured using a gas-exchange technique applied to branches of trees. The extent of spectral changes in incident solar radiation was assessed using a spectroradiometer.We found significantly higher NEE (up to 150%) during the cloudy periods compared with the sunny periods at corresponding PPFDs. Prevailing diffuse radiation under the cloudy days resulted in a significantly lower compensation irradiance (by ca. 50% and 70%), while apparent quantum yield was slightly higher (by ca. 7%) at canopy level and significantly higher (by ca. 530%) in sun-acclimated shoots. The main reasons for these differences appear to be (1) more favourable microclimatic conditions during cloudy periods, (2) stimulation of photochemical reactions and stomatal opening via an increase of blue/red light ratio, and (3) increased penetration of light into the canopy and thus a more equitable distribution of light between leaves.Our analyses identified the most important reason of enhanced NEE under cloudy sky conditions to be the effective penetration of diffuse radiation to lower depths of the canopy. This subsequently led to the significantly higher solar equivalent leaf area compared with the direct radiation. Most of the leaves in such dense canopy are in deep shade, with marginal or negative carbon balances during sunny days. These findings show that the energy of diffuse, compared with direct, solar radiation is used more efficiently in assimilation processes at both leaf and canopy levels.

Boreal forests comprise 73% of the world’s coniferous forests. Based on forest floor measurements, they have been considered a significant natural sink of methane (CH4) and a natural source of nitrous oxide (N2O), both of which are important greenhouse gases. However, the role of trees, especially conifers, in ecosystem N2O and CH4 exchange is only poorly understood. We show for the first time that mature Scots pine (Pinus sylvestris L.) trees consistently emit N2O and CH4 from both stems and shoots. The shoot fluxes of N2O and CH4 exceeded the stem flux rates by 16 and 41 times, respectively. Moreover, higher stem N2O and CH4 fluxes were observed from wet than from dry areas of the forest. The N2O release from boreal pine forests may thus be underestimated and the uptake of CH4 may be overestimated when ecosystem flux calculations are based solely on forest floor measurements. The contribution of pine trees to the N2O and CH4 exchange of the boreal pine forest seems to increase considerably under high soil water content, thus highlighting the urgent need to include tree-emissions in greenhouse gas emission inventories.

No abstract

Shoots and roots are autotrophic and heterotrophic organs of plants with different physiological functions. Do they have different metabolomes? Do their metabolisms respond differently to environmental changes such as drought? We used metabolomics and elemental analyses to answer these questions. First, we show that shoots and roots have different metabolomes and nutrient and elemental stoichiometries. Second, we show that the shoot metabolome is much more variable among species and seasons than is the root metabolome. Third, we show that the metabolic response of shoots to drought contrasts with that of roots; shoots decrease their growth metabolism (lower concentrations of sugars, amino acids, nucleosides, N, P, and K), and roots increase it in a mirrored response. Shoots are metabolically deactivated during drought to reduce the consumption of water and nutrients, whereas roots are metabolically activated to enhance the uptake of water and nutrients, together buffering the effects of drought, at least at the short term.

Root exudates comprise a large variety of compounds released by plants into the rhizosphere, including low-molecular-weight primary metabolites (particularly saccharides, amino acids and organic acids) and secondary metabolites (phenolics, flavonoids and terpenoids). Changes in exudate composition could have impacts on the plant itself, on other plants, on soil properties (e.g. amount of soil organic matter), and on soil organisms. The effects of drought on the composition of root exudates, however, have been rarely studied. We used an ecometabolomics approach to identify the compounds in the exudates of Quercus ilex (holm oak) under an experimental drought gradient and subsequent recovery. Increasing drought stress strongly affected the composition of the exudate metabolome. Plant exudates under drought consisted mainly of secondary metabolites (71% of total metabolites) associated with plant responses to drought stress, whereas the metabolite composition under recovery shifted towards a dominance of primary metabolites (81% of total metabolites). These results strongly suggested that roots exude the most abundant root metabolites. The exudates were changed irreversibly by the lack of water under extreme drought conditions, and the plants could not recover.

Summary1. Cloud cover affects carbon exchange between biota and the atmosphere. Recent studies have demonstrated that an increase in the diffuse radiation fraction enhances the photosynthetic efficiency of canopies. Although the exact mechanism behind this effect is not clear, a more even distribution of light among leaves across the vertical profile of the canopy is considered to be the most important cause of this difference. 2. To test this hypothesis, the net ecosystem production (NEP) of a Norway spruce forest (30-year-old) was measured under cloudy and sunny skies by the eddy covariance method. In parallel, measurements of the diurnal courses of gas exchange and chlorophyll fluorescence parameters were made in the upper sun (5th whorl; 1-year-old needles), middle (8th and 10th whorl; 1-and 2-year-old needles) and lower shade (15th whorl; >2-year-old needles) shoots. 3. The higher diffuse radiation fraction during cloudy days resulted in significantly higher ecosystem carbon uptake than at corresponding incident photosynthetic photon flux density on sunny days. Our shoot-level data show that shoots from deep within the canopy contribute substantially to the overall carbon balance during cloudy days. But, although shade-adapted shoots had a markedly positive carbon balance over a 24-h period on cloudy days, their performance was impaired on sunny days contributing only a marginal or even negative carbon balance from the middle and shaded parts of the canopy. The uppermost sun shoots contributed 78% of the total carbon assimilated during a sunny day, but only 43% during a cloudy day. 4. In addition, afternoon depression of canopy NEP and CO 2 assimilation rates of the uppermost shoots (5th and 8th whorl) occurred in response to irradiance on sunny days, characterized by significant decreases in CO 2 uptake and apparent quantum yield; however, this depression did not occur under cloudy conditions. Stomatal and non-stomatal regulations of carbon assimilation in the afternoon are discussed.