The contribution of photodegradation to litter decomposition in a temperate forest gap and understorey.

Litter decomposition determines carbon (C) backflow to the atmosphere and ecosystem nutrient cycling. Although sunlight provides the indispensable energy for terrestrial biogeochemical processes, the role of photodegradation in decomposition has been relatively neglected in productive mesic ecosystems. To quantify the effects of this variation, we conducted a factorial experiment in the understorey of a temperate deciduous forest and an adjacent gap, using spectral-attenuation-filter treatments. Exposure to the full spectrum of sunlight increased decay rates by nearly 120% and the effect of blue light contributed 75% of this increase. Scaled-up to the whole forest ecosystem, this translates to 13% loss of leaf-litter C through photodegradation over the year of our study for a scenario of 20% gap. Irrespective of the spectral composition, herbaceous and shrub litter lost mass faster than tree litter, with photodegradation contributing the most to surface litter decomposition in forest canopy gaps. Across species, the initial litter lignin and polyphenolic contents predicted photodegradation by blue light and ultraviolet B (UV-B) radiation, respectively. We concluded that photodegradation, modulated by litter quality, is an important driver of decomposition, not just in arid areas, but also in mesic ecosystems such as temperate deciduous forests following gap opening.

A perspective on ecologically relevant plant-UV research and its practical application.

Plants perceive ultraviolet-B (UV-B) radiation through the UV-B photoreceptor UV RESISTANCE LOCUS 8 (UVR8), and initiate regulatory responses via associated signalling networks, gene expression and metabolic pathways. Various regulatory adaptations to UV-B radiation enable plants to harvest information about fluctuations in UV-B irradiance and spectral composition in natural environments, and to defend themselves against UV-B exposure. Given that UVR8 is present across plant organs and tissues, knowledge of the systemic signalling involved in its activation and function throughout the plant is important for understanding the context of specific responses. Fine-scale understanding of both UV-B irradiance and perception within tissues and cells requires improved application of knowledge about UV-attenuation in leaves and canopies, warranting greater consideration when designing experiments. In this context, reciprocal crosstalk among photoreceptor-induced pathways also needs to be considered, as this appears to produce particularly complex patterns of physiological and morphological response. Through crosstalk, plant responses to UV-B radiation go beyond simply UV-protection or amelioration of damage, but may give cross-protection over a suite of environmental stressors. Overall, there is emerging knowledge showing how information captured by UVR8 is used to regulate molecular and physiological processes, although understanding of upscaling to higher levels of organisation, i.e. organisms, canopies and communities remains poor. Achieving this will require further studies using model plant species beyond Arabidopsis, and that represent a broad range of functional types. More attention should also be given to plants in natural environments in all their complexity, as such studies are needed to acquire an improved understanding of the impact of climate change in the context of plant-UV responses. Furthermore, broadening the scope of experiments into the regulation of plant-UV responses will facilitate the application of UV radiation in commercial plant production. By considering the progress made in plant-UV research, this perspective highlights prescient topics in plant-UV photobiology where future research efforts can profitably be focussed. This perspective also emphasises burgeoning interdisciplinary links that will assist in understanding of UV-B effects across organisational scales and gaps in knowledge that need to be filled so as to achieve an integrated vision of plant responses to UV-radiation.

Solar UV-A radiation and blue light enhance tree leaf litter decomposition in a temperate forest.

Sunlight can accelerate the decomposition process through an ensemble of direct and indirect processes known as photodegradation. Although photodegradation is widely studied in arid environments, there have been few studies in temperate regions. This experiment investigated how exposure to solar radiation, and specifically UV-B, UV-A, and blue light, affects leaf litter decomposition under a temperate forest canopy in France. For this purpose, we employed custom-made litterbags built using filters that attenuated different regions of the solar spectrum. Litter mass loss and carbon to nitrogen (C:N) ratio of three species: European ash (Fraxinus excelsior), European beech (Fagus sylvatica) and pedunculate oak (Quercus robur), differing in their leaf traits and decomposition rate, were analysed over a period of 7-10 months. Over the entire period, the effect of treatments attenuating blue light and solar UV radiation on leaf litter decomposition was similar to that of our dark treatment, where litter lost 20-30% less mass and had a lower C:N ratio than under the full-spectrum treatment. Moreover, decomposition was affected more by the filter treatment than mesh size, which controlled access by mesofauna. The effect of filter treatment differed among the three species and appeared to depend on litter quality (and especially C:N), producing the greatest effect in recalcitrant litter (F. sylvatica). Even under the reduced irradiance found in the understorey of a temperate forest, UV radiation and blue light remain important in accelerating surface litter decomposition.

Canopy structure and phenology modulate the impacts of solar radiation on C and N dynamics during litter decomposition in a temperate forest.

Decomposition of plant organic matter plays a key role in the terrestrial biogeochemical cycles. Sunlight has recently been identified as an important contributor to carbon [C] turnover through photodegradation, accelerating decomposition even in forest ecosystems where understorey solar irradiance remains relatively low. However, it is uncertain how C and nutrients dynamics respond to fluctuations in solar spectral irradiance caused by canopy structure (understorey vs. gaps) and season (open vs. closed canopy phenology). Spectral-attenuation treatments were used to compare litter decomposition over eight months, covering canopy phenology, in a temperate deciduous forest and an adjacent gap. Exposure to the full spectrum of sunlight increased the loss of litter C and lignin by 75% and 64% in the forest gap, and blue light was responsible for respectively 27% and 42% of that loss. Whereas in the understorey, C and lignin loss were similar among spectral-attenuation treatments over the experimental period, except prior to and during spring canopy flush when exposure to the full spectrum of sunlight promoted C loss by 15% overall, 80% of which was attributable to ultraviolet-B (UV-B) radiation. Nitrogen [N] was immobilized in the understorey during canopy flush before the canopy completely closed but N was swiftly released during canopy leaf-fall. Our study suggests that blue-driven photodegradation plays an important role in lignin decomposition and N dynamics in canopy gaps, whereas seasonal canopy phenology affecting sunlight reaching the forest floor drastically changes patterns of C and N in litter during decomposition. Hence, including sunlight dynamics driven by canopy structure and phenology would improve estimates of biogeochemical cycling in forests responding to changes in climate and land-use.

Ultraviolet radiation accelerates photodegradation under controlled conditions but slows the decomposition of senescent leaves from forest stands in southern Finland.

Depending on the environment, sunlight can positively or negatively affect litter decomposition, through the ensemble of direct and indirect processes constituting photodegradation. Which of these processes predominate depends on the ecosystem studied and on the spectral composition of sunlight received. To examine the relevance of photodegradation for litter decomposition in forest understoreys, we filtered ultraviolet radiation (UV) and blue light from leaves of Fagus sylvatica and Betula pendula at two different stages of senescence in both a controlled-environment experiment and outdoors in four different forest stands (Picea abies, Fagus sylvatica, Acer platanoides, Betula pendula). Controlling for leaf orientation and initial differences in leaf chlorophyll and flavonol concentrations; we measured mass loss at the end of each experiment and characterised the phenolic profile of the leaf litter following photodegradation. In most forest stands, less mass was lost from decomposing leaves that received solar UV radiation compared with those under UV-attenuating filters, while in the controlled environment UV-A radiation either slightly accelerated or had no significant effect on photodegradation, according to species identity. Only a few individual phenolic compounds were affected by our different filter treatments, but photodegradation did affect the phenolic profile. We can conclude that photodegradation has a small stand- and species-specific effect on the decomposition of surface leaf litter in forest understoreys during the winter following leaf fall in southern Finland. Photodegradation was wavelength-dependent and modulated by the canopy species filtering sunlight and likely creating different combinations of spectral composition, moisture, temperature and snowpack characteristics.

Seasonal Patterns in Spectral Irradiance and Leaf UV-A Absorbance Under Forest Canopies.

Plants commonly respond to UV radiation through the accumulation of flavonoids and related phenolic compounds which potentially ameliorate UV-damage to crucial internal structures. However, the seasonal dynamics of leaf flavonoids corresponding to epidermal UV absorbance is highly variable in nature, and it remains uncertain how environmental factors combine to govern flavonoid accumulation and degradation. We studied leaf UV-A absorbance of species composing the understorey plant community throughout two growing seasons under five adjacent tree canopies in southern Finland. We compared the relationship between leaf flavonol index (Iflav-repeatedly measured with an optical leaf clip Dualex) and measured spectral irradiance, understorey and canopy phenology, air temperature and snowpack variables, whole leaf flavonoid extracts, and leaf age. Strong seasonal patterns and stand-related differences were apparent in Iflav of both understorey plant communities and individual species, including divergent trends in Iflav during spring and autumn. Comparing the heterogeneity of the understorey light environment and its spectral composition in looking for potential drivers of seasonal changes in Iflav, we found that unweighted UV-A irradiance, or the effective UV dose calculated according to the biological spectral weighting function (BSWF) for plant growth (PG action spectrum), in understorey shade had a strong relationship with Iflav. Furthermore, understorey species seemed to adjust Iflav to low background diffuse irradiance rather than infrequent high direct-beam irradiance in sunflecks during summer, since leaves produced during or after canopy closure had low Iflav. In conclusion, we found the level of epidermal flavonoids in forest understorey species to be plastic, adjusting to climatic conditions, and differing according to species' leaf retention strategy and new leaf production, all of which contribute to the seasonal trends in leaf flavonoids found within forest stands.

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.