Development of a sink-source interaction model for the growth of short-rotation coppice willow and in silico exploration of genotype×environment effects.

Identifying key performance traits is essential for elucidating crop growth processes and breeding. In Salix spp., genotypic diversity is being exploited to tailor new varieties to overcome environmental yield constraints. Process-based models can assist these efforts by identifying key parameters of yield formation for different genotype×environment (G×E) combinations. Here, four commercial willow varieties grown in contrasting environments (west and south-east UK) were intensively sampled for growth traits over two 2-year rotations. A sink-source interaction model was developed to parameterize the balance of source (carbon capture/mobilization) and sink formation (morphogenesis, carbon allocation) during growth. Global sensitivity analysis consistently identified day length for the onset of stem elongation as most important factor for yield formation, followed by various 'sink>source' controlling parameters. In coastal climates, the chilling control of budburst ranked higher compared with the more eastern climate. Sensitivity to drought, including canopy size and rooting depth, was potentially growth limiting in the south-east and west of the UK. Potential yields increased from the first to the second rotation, but less so for broad- than for narrow-leaved varieties (20 and 47%, respectively), which had established less well initially (-19%). The establishment was confounded by drought during the first rotation, affecting broad- more than narrow-leaved canopy phenotypes (-29%). The analysis emphasized quantum efficiency at low light intensity as key to assimilation; however, on average, sink parameters were more important than source parameters. The G×E pairings described with this new process model will help to identify parameters of sink-source control for future willow breeding.

Climate change alters leaf anatomy, but has no effects on volatile emissions from Arctic plants.

Biogenic volatile organic compound (BVOC) emissions are expected to change substantially because of the rapid advancement of climate change in the Arctic. BVOC emission changes can feed back both positively and negatively on climate warming. We investigated the effects of elevated temperature and shading on BVOC emissions from arctic plant species Empetrum hermaphroditum, Cassiope tetragona, Betula nana and Salix arctica. Measurements were performed in situ in long-term field experiments in subarctic and high Arctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. In order to assess whether the treatments had resulted in anatomical adaptations, we additionally examined leaf anatomy using light microscopy and scanning electron microscopy. Against expectations based on the known temperature and light-dependency of BVOC emissions, the emissions were barely affected by the treatments. In contrast, leaf anatomy of the studied plants was significantly altered in response to the treatments, and these responses appear to differ from species found at lower latitudes. We suggest that leaf anatomical acclimation may partially explain the lacking treatment effects on BVOC emissions at plant shoot-level. However, more studies are needed to unravel why BVOC emission responses in arctic plants differ from temperate species.

Photoinhibition-like damage to the photosynthetic apparatus in plant leaves induced by submergence treatment in the dark.

Submergence is a common type of environmental stress for plants. It hampers survival and decreases crop yield, mainly by inhibiting plant photosynthesis. The inhibition of photosynthesis and photochemical efficiency by submergence is primarily due to leaf senescence and excess excitation energy, caused by signals from hypoxic roots and inhibition of gas exchange, respectively. However, the influence of mere leaf-submergence on the photosynthetic apparatus is currently unknown. Therefore, we studied the photosynthetic apparatus in detached leaves from four plant species under dark-submergence treatment (DST), without influence from roots and light. Results showed that the donor and acceptor sides, the reaction center of photosystem II (PSII) and photosystem I (PSI) in leaves were significantly damaged after 36 h of DST. This is a photoinhibition-like phenomenon similar to the photoinhibition induced by high light, as further indicated by the degradation of PsaA and D1, the core proteins of PSI and PSII. In contrast to previous research, the chlorophyll content remained unchanged and the H2O2 concentration did not increase in the leaves, implying that the damage to the photosynthetic apparatus was not caused by senescence or over-accumulation of reactive oxygen species (ROS). DST-induced damage to the photosynthetic apparatus was aggravated by increasing treatment temperature. This type of damage also occurred in the anaerobic environment (N2) without water, and could be eliminated or restored by supplying air to the water during or after DST. Our results demonstrate that DST-induced damage was caused by the hypoxic environment. The mechanism by which DST induces the photoinhibition-like damage is discussed below.

Combined enhancements of temperature and UVB influence growth and phenolics in clones of the sexually dimorphic Salix myrsinifolia.

Although several climatic factors are expected to change simultaneously in the future, the effect of such combined changes on plants have seldom been tested under field conditions. We report on a field experiment with dark-leaved willow, Salix myrsinifolia, subjected to enhancements in ultraviolet-A (UVA), UVB radiation and temperature, setup in Joensuu, Eastern Finland. S. myrsinifolia is a dioecious species, known as an important food plant for many herbivores. Cuttings of eight clones, four of each sex, of dark-leaved willow were planted in the field in spring 2009. In both 2009 and 2010, the total biomass increased significantly with temperature, and in 2010 there was an additive effect of UVB radiation. Both height and diameter increased with temperature in 2009, while the effect on height growth ceased in 2010. Males had greater diameter growth than females in 2010. Most phenolic compounds in the leaves decreased under enhanced temperature in both growing seasons. In 2010, four of six salicylates increased in response to enhanced temperature. Some quercetin derivatives increased under enhanced UVB radiation. Females had higher concentrations of chlorogenic acids than males, and while enhanced temperature reduced chlorogenic acid in females only, luteolins were reduced only in males. In summary, the combined enhancements gave no effects in addition to those that appeared under the single-factor treatments, except for the additive effect of UVB on temperature-increased biomass. The few gender-related differences found in response to climate change do not allow any marked expectations of future climate-induced changes in sex ratios.

Dark-leaved willow (Salix myrsinifolia) is resistant to three-factor (elevated CO2, temperature and UV-B-radiation) climate change.

Elevated carbon dioxide (CO2 ), temperature (T) and ultraviolet-B (UV-B) radiation may affect plant growth and secondary chemistry in different directions, but the effect of the combination of the three factors has seldom been tested. Here, we grew four dark-leaved willow (Salix myrsinifolia) clones under combinations of ambient or elevated CO2, T and UV-B radiation in top-closed chambers for 7 wk. Elevated UV-B had no effects on growth or phenolic compounds, and there were no significant interactions between UV-B, CO2 and T. CO2 alone increased most growth parameters, but the magnitude of the effect varied among the clones. Total phenolics increased at elevated CO2 , whereas they decreased at elevated T. The responses varied between the clones. The results imply that dark-leaved willow are fairly resistant to the applied three-factor climate change, probably because of high constitutive defense. However, the interactions between clone and climate change factors implies that some clones are more susceptible than the species as a whole.

Improved UV-B screening capacity does not prevent negative effects of ambient UV irradiance on PSII performance in High Arctic plants. Results from a six year UV exclusion study.

Long-term responses of ambient solar ultraviolet (UV) radiation were investigated on Salix arctica and Vaccinium uliginosum in a High Arctic heath ecosystem in Zackenberg, northeast Greenland. Over a period of six years, UV exclusion was conducted in the growing season by means of filters: 60% UV-B reduction, 90% UV-B+UV-A reduction, UV transparent filter control, and an open control without filter. Plant responses were evaluated using specific leaf area, leaf content of UV-B absorbing compounds and PSII performance parameters derived from chlorophyll-a fluorescence induction curves. Based on the JIP-test, we calculated the total performance index PI(total), which includes the integrating antennae, the PSII reaction center, intersystem electron transport and reduction of PSI end acceptors-dependent parameters. In both species, UV exclusion significantly decreased the content of UV-B-absorbing compounds. Salix increased its specific leaf area, while Vaccinium decreased it. UV exclusion increased the PI(total) in both species during all six years of experimentation. This response was governed by a significantly decreased RC/ABS, a marginally non-significant increased ET(o)/TR(o) and a significantly increased TR(o)/ABS=F(V)/F(M) and RE(o)/ET(o). These results demonstrate the current level of ambient UV-B to decrease PSII performance significantly in these High Arctic plants. It appears that the two plant species both have improved their UV-screening capacity, but through different strategies, although this did not sufficiently prevent negative effects of the ambient UV radiation. We argue the decreased PSII performance to be part of a response decreasing plant carbon uptake. We speculate the negative effects on PSII performance mediated by ambient UV irradiance to be present in years where warming induces early snowmelt, exposing the vegetation to high spring UV-B, and to be present in the future to the degree the ozone layer is not fully recovered.

Effects of photooxidation on membrane integrity in Salix nigra seeds.

BACKGROUND AND AIMS: Salix nigra seeds are desiccation-tolerant, as are orthodox seeds, although in contrast to other orthodox seeds they lose viability in a few weeks at room temperature. They also differ in that the chloroplasts of the embryo tissues conserve their chlorophyll and endomembranes. The aim of this paper was to investigate the role of chlorophyll in seed deterioration. METHODS: Seeds were aged at different light intensities and atmospheric conditions. Mean germination time and normal and total germination were evaluated. The formation of free radicals was assessed using electronic spin resonance spectroscopy, and changes in the fatty acid composition from phospholipids, galactolipids and triglycerides using gas-liquid chromatography. Membrane integrity was studied with electronic spin resonance spin probe techniques, electrolyte leakage and transmission electron microscopy. KEY RESULTS: Light and oxygen played an important role in free-radical generation, causing a decrease in normal germination and an increase in mean germination time. Both indices were associated with a decrease in polyunsaturated fatty acids derived from membrane lipids as phospholipids and galactolipids. The detection of damage in thylakoid membranes and an increase in plasmalemma permeability were consistent with the decrease in both types of lipids. Triglycerides remained unchanged. Light-induced damage began in outermost tissues and spread inwards, decreasing normal germination. CONCLUSIONS: Salix nigra seeds were very susceptible to photooxidation. The thylakoid membranes appeared to be the first target of the photooxidative process since there were large decreases in galactolipids and both these lipids and the activated chlorophyll are contiguous in the structure of that membrane. Changes in normal germination and mean germination time could be explained by the deteriorative effects of oxidation.

Evapotranspiration and crop coefficient of poplar and willow short-rotation coppice used as vegetation filter.

Ten-day evapotranspiration (ETc) and crop coefficient (k(c)) of willow and poplar SRC used as vegetation filter and grown under fertilised (F) and unfertilised (NF) conditions, were determined for two successive growing seasons using volumetric lysimeters. During the first growing season, total ETc observed was, respectively, 620 (NF)-1190 (F)mm in willow and 590 (NF)-725 (F) in poplar. During the second growing season, ETc showed a general increase, mainly in fertilised lysimeters where it ranged between 890 (NF)-1790 mm (F) in willow and 710 (NF)-1100 mm (NF) in poplar. kc reached in both years its maximum between the end of August and the beginning of September. In 2004 maximum kc ranged from 1.25-2.84 in willow and 1.06-1.90 in poplar, whereas in 2005 it ranged from 1.97-5.30 in willow and 1.71-4.28 in poplar. ETc seemed to be strongly correlated to plant development and mainly dependent on its nutritional status rather than on the differences between the species.