Understorey light quality affects leaf pigments and leaf phenology in different plant functional types.

Forest understorey plants receive most sunlight in springtime before canopy closure, and in autumn following leaf-fall. We hypothesised that plant species must adjust their phenological and photoprotective strategies in response to large changes in the spectral composition of the sunlight they receive. Here, we identified how plant species growing in northern deciduous and evergreen forest understoreys differ in their response to blue light and ultraviolet (UV) radiation according to their functional strategy. We installed filters in a forest understorey in southern Finland, to create the following treatments attenuating: UV radiation below 350 nm, all UV radiation (< 400 nm), all blue light and UV radiation (< 500 nm), and a transparent control. In eight species, representing different functional strategies, we assessed leaf optical properties, phenology, and epidermal flavonoid contents over two years. Blue light accelerated leaf senescence in all species measured in the understorey, apart from Quercus robur seedlings, whereas UV radiation only accelerated leaf senescence in Acer platanoides seedlings. More light-demanding species accumulated flavonols in response to seasonal changes in light quality compared to shade-tolerant and wintergreen species and were particularly responsive to blue light. Reduction of blue and UV radiation under shade reveals an important role for microclimatic effects on autumn phenology and leaf photoprotection. An extension of canopy cover under climate change, and its associated suppression of understorey blue light and UV radiation, may delay leaf senescence for understorey species with an autumn niche.

The roles of species' relatedness and climate of origin in determining optical leaf traits over a large set of taxa growing at high elevation and high latitude.

Climate change is driving many mountain plant species to higher elevations and northern plant species to higher latitudes. However, various biotic or abiotic constraints may restrict any range shift, and one relevant factor for migration to higher elevations could be species' ability to tolerate high UV-doses. Flavonoids are engaged in photoprotection, but also serve multiple ecological roles. We compared plant optical leaf trait responses of a large set of taxa growing in two botanical gardens (French Alps and southern Finland), considering potential constraints imposed by the relatedness of taxa and the legacy of climatic conditions at plants' original collection sites. The segregation of optically measured leaf traits along the phylogeny was studied using a published mega-tree GBOTB.extended.tre for vascular plants as a backbone. For a subset of taxa, we investigated the relationship between climatic conditions (namely solar radiation, temperature and precipitation at a coarse scale) at the plants' original collection site and current trait values. Upon testing the phylogenetic signal (Pagel's λ), we found a significant difference but intermediate lambda values overall for flavonol or flavone index (Iflav) and anthocyanin index (Iant), indicating that phylogenetic relatedness alone failed to explain the changes in trait values under a Brownian motion model of trait evolution. The local analysis (local indicator of phylogenetic association) indicated mostly positive autocorrelations for Iflav i.e. similarities in optically measured leaf traits, often among species from the same genus. We found significant relationships between climatic variables and leaf chlorophyll index (Ichl), but not Iflav, particularly for annual solar radiation. Changes in plants' Iflav across microhabitats differing in UV irradiance and predominately high F v /F m indicated that most plants studied had sufficient flexibility in photoprotection, conferred by Iflav, to acclimate to contemporary UV irradiances in their environment. While not explaining the mechanisms behind observed trait values, our findings do suggest that some high-elevation taxa display similar leaf flavonoid accumulation responses. These may be phylogenetically constrained and hence moderate plants' capacity to adjust to new combinations of environmental conditions resulting from climate change.

Do UV-A radiation and blue light during growth prime leaves to cope with acute high light in photoreceptor mutants of Arabidopsis thaliana?

We studied how plants acclimated to growing conditions that included combinations of blue light (BL) and ultraviolet (UV)-A radiation, and whether their growing environment affected their photosynthetic capacity during and after a brief period of acute high light (as might happen during an under-canopy sunfleck). Arabidopsis thaliana Landsberg erecta wild-type were compared with mutants lacking functional blue light and UV photoreceptors: phototropin 1, cryptochromes (CRY1 and CRY2) and UV RESISTANT LOCUS 8 (uvr8). This was achieved using light-emitting-diode (LED) lamps in a controlled environment to create treatments with or without BL, in a split-plot design with or without UV-A radiation. We compared the accumulation of phenolic compounds under growth conditions and after exposure to 30 min of high light at the end of the experiment (46 days), and likewise measured the operational efficiency of photosystem II (ϕPSII, a proxy for photosynthetic performance) and dark-adapted maximum quantum yield (Fv /Fm to assess PSII damage). Our results indicate that cryptochromes are the main photoreceptors regulating phenolic compound accumulation in response to BL and UV-A radiation, and a lack of functional cryptochromes impairs photosynthetic performance under high light. Our findings also reveal a role for UVR8 in accumulating flavonoids in response to a low UV-A dose. Interestingly, phototropin 1 partially mediated constitutive accumulation of phenolic compounds in the absence of BL. Low-irradiance BL and UV-A did not improve ϕPSII and Fv /Fm upon our acute high-light treatment; however, CRYs played an important role in ameliorating high-light stress.

UV-screening and springtime recovery of photosynthetic capacity in leaves of Vaccinium vitis-idaea above and below the snow pack.

Evergreen plants in boreal biomes undergo seasonal hardening and dehardening adjusting their photosynthetic capacity and photoprotection; acclimating to seasonal changes in temperature and irradiance. Leaf epidermal ultraviolet (UV)-screening by flavonols responds to solar radiation, perceived in part through increased ultraviolet-B (UV-B) radiation, and is a candidate trait to provide cross-photoprotection. At Hyytiälä Forestry Station, central Finland, we examined whether the accumulation of flavonols was higher in leaves of Vaccinium vitis-idaea L. growing above the snowpack compared with those below the snowpack. We found that leaves exposed to colder temperatures and higher solar radiation towards the top of hummocks suffered greater photoinhibition than those at the base of hummocks. Epidermal UV-screening was highest in upper-hummock leaves, particularly during winter when lower leaves were beneath the snowpack. There was also a negative relationship between indices of flavonols and anthocyanins across all leaves suggesting fine-tuning of flavonoid composition for screening vs. antioxidant activity in response to temperature and irradiance. However, the positive correlation between the maximum quantum yield of photosystem II photochemistry (Fv/Fm) and flavonol accumulation in upper hummock leaves during dehardening did not confer on them any greater cross-protection than would be expected from the general relationship of Fv/Fm with temperature and irradiance (throughout the hummocks). Irrespective of timing of snow-melt, photosynthesis fully recovered in all leaves, suggesting that V. vitis-idaea has the potential to exploit the continuing trend for longer growing seasons in central Finland without incurring significant impairment from reduced duration of snow cover.

Assessing scale-wise similarity of curves with a thick pen: As illustrated through comparisons of spectral irradiance.

Forest canopies create dynamic light environments in their understorey, where spectral composition changes among patterns of shade and sunflecks, and through the seasons with canopy phenology and sun angle. Plants use spectral composition as a cue to adjust their growth strategy for optimal resource use. Quantifying the ever-changing nature of the understorey light environment is technically challenging with respect to data collection. Thus, to capture the simultaneous variation occurring in multiple regions of the solar spectrum, we recorded spectral irradiance from forest understoreys over the wavelength range 300-800 nm using an array spectroradiometer. It is also methodologically challenging to analyze solar spectra because of their multi-scale nature and multivariate lay-out. To compare spectra, we therefore used a novel method termed thick pen transform (TPT), which is simple and visually interpretable. This enabled us to show that sunlight position in the forest understorey (i.e., shade, semi-shade, or sunfleck) was the most important factor in determining shape similarity of spectral irradiance. Likewise, the contributions of stand identity and time of year could be distinguished. Spectra from sunflecks were consistently the most similar, irrespective of differences in global irradiance. On average, the degree of cross-dependence increased with increasing scale, sometimes shifting from negative (dissimilar) to positive (similar) values. We conclude that the interplay of sunlight position, stand identity, and date cannot be ignored when quantifying and comparing spectral composition in forest understoreys. Technological advances mean that array spectroradiometers, which can record spectra contiguously over very short time intervals, are being widely adopted, not only to measure irradiance under pollution, clouds, atmospheric changes, and in biological systems, but also spectral changes at small scales in the photonics industry. We consider that TPT is an applicable method for spectral analysis in any field and can be a useful tool to analyze large datasets in general.

How does solar ultraviolet-B radiation improve drought tolerance of silver birch (Betula pendula†Roth.) seedlings?

We hypothesized that solar ultraviolet (UV) radiation would protect silver birch seedlings from the detrimental effects of water stress through a coordinated suite of trait responses, including morphological acclimation, improved control of water loss through gas exchange and hydraulic sufficiency. To better understand how this synergetic interaction works, plants were grown in an experiment under nine treatment combinations attenuating ultraviolet-A and ultraviolet-B (UVB) from solar radiation together with differential watering to create water-deficit conditions. In seedlings under water deficit, UV attenuation reduced height growth, leaf production and leaf length compared with seedlings receiving the full spectrum of solar radiation, whereas the growth and morphology of well-watered seedlings was largely unaffected by UV attenuation. There was an interactive effect of the treatment combination on water relations, which was more apparent as a change in the water potential at which leaves wilted or plants died than through differences in gas exchange. This suggests that changes occur in the cell wall elastic modulus or accumulation of osmolites in cells under UVB. Overall, the strong negative effects of water deficit are partially ameliorated by solar UV radiation, whereas well-watered silver birch seedlings are slightly disadvantaged by the solar UV radiation they receive.