Giant sequoia (Sequoiadendron giganteum) in the UK: carbon storage potential and growth rates.

Giant sequoias (Sequoiadendron giganteum) are some of the UK's largest trees, despite only being introduced in the mid-nineteenth century. There are an estimated half a million giant sequoias and closely related coastal redwoods (Sequoia sempervirens) in the UK. Given the recent interest in planting more trees, partly due to their carbon sequestration potential and also their undoubted public appeal, an understanding of their growth capability is important. However, little is known about their growth and carbon uptake under UK conditions. Here, we focus on S. giganteum and use three-dimensional terrestrial laser scanning to perform detailed structural measurements of 97 individuals at three sites covering a range of different conditions, to estimate aboveground biomass (AGB) and annual biomass accumulation rates. We show that UK-grown S. giganteum can sequester carbon at a rate of 85 kg yr-1, varying with climate, management and age. We develop new UK-specific allometric models for S. giganteum that fit the observed AGB with r 2 > 0.93 and bias < 2% and can be used to estimate S. giganteum biomass more generally. This study provides the first estimate of the growth and carbon sequestration of UK open-grown S. giganteum and provides a baseline for estimating their longer-term carbon sequestration capacity.

Radiation and temperature drive diurnal variation of aerobic methane emissions from Scots pine canopy.

Methane emissions from plant foliage may play an important role in the global methane cycle, but their size and the underlying source processes remain poorly understood. Here, we quantify methane fluxes from the shoots of Scots pine trees, a dominant tree species in boreal forests, to identify source processes and environmental drivers, and we evaluate whether these fluxes can be constrained at the ecosystem-level by eddy covariance flux measurements. We show that shoot-level measurements conducted in forest, garden, or greenhouse settings; on mature trees and saplings; manually and with an automated CO2-, temperature-, and water-controlled chamber system; and with multiple methane analyzers all resulted in comparable daytime fluxes (0.144 ± 0.019 to 0.375 ± 0.074 nmol CH4 g-1 foliar d.w. h-1). We further find that these emissions exhibit a pronounced diurnal cycle that closely follows photosynthetically active radiation and is further modulated by temperature. These diurnal patterns indicate that methane production is associated with diurnal cycle of sunlight, indicating that this production is either a byproduct of photosynthesis-associated biochemical reactions (e.g., the methionine cycle) or produced through nonenzymatic photochemical reactions in plant biomass. Moreover, we identified a light-dependent component in stand-level methane fluxes, which showed order-of-magnitude agreement with shoot-level measurements (0.968 ± 0.031 nmol CH4 g-1 h-1) and which provides an upper limit for shoot methane emissions.

Plasma membrane aquaporins of the PIP1 and PIP2 subfamilies facilitate hydrogen peroxide diffusion into plant roots.

BACKGROUND: The permeability of plasma membrane aquaporins (PIPs) to small solutes other than water greatly diversifies their potential functions in plant development and metabolic processes. One such process is stress signalling in which hydrogen peroxide (H2O2) plays a major role. Based on transport assays carried out in yeast, there are differences in the degree to which PIPs of Arabidopsis thaliana, are permeable to H2O2 and thus they may differentially facilitate transmembrane diffusion. Here, we test whether specific PIPs aid in the transmembrane diffusion of H2O2 to such an extent that knocking-out PIPs affects plant phenotype. We examined changes in growth and morphology, including biomass accumulation, root system architecture and relative water content, as well as gas exchange, across two H2O2 treatments in knockout mutants of A. thaliana. RESULTS: We could infer that PIP-type aquaporins are permeable to H2O2 in planta and that this permeability is physiologically relevant in a plant's response to oxidative stress. In particular, the lack of functional PIP2;3 confers resistance to exogenously applied H2O2 indicating that it facilitates H2O2 entry into root cells. Additionally, PIP1;1 and PIP2;6 were found to facilitate H2O2 diffusion, while PIP2;2 is required for proper root growth under controlled conditions. MAIN FINDINGS: We conclude that PIPs are physiologically relevant conduits for H2O2 diffusion in the A. thaliana roots and participate in the regulation of stress responses.

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.

The success of the Montreal Protocol in mitigating interactive effects of stratospheric ozone depletion and climate change on the environment.

The Montreal Protocol and its Amendments have been highly effective in protecting the stratospheric ozone layer, preventing global increases in solar ultraviolet-B radiation (UV-B; 280-315 nm) at Earth's surface, and reducing global warming. While ongoing and projected changes in UV-B radiation and climate still pose a threat to human health, food security, air and water quality, terrestrial and aquatic ecosystems, and construction materials and fabrics, the Montreal Protocol continues to play a critical role in protecting Earth's inhabitants and ecosystems by addressing many of the United Nations Sustainable Development Goals.

Chlorophyll a fluorescence illuminates a path connecting plant molecular biology to Earth-system science.

For decades, the dynamic nature of chlorophyll a fluorescence (ChlaF) has provided insight into the biophysics and ecophysiology of the light reactions of photosynthesis from the subcellular to leaf scales. Recent advances in remote sensing methods enable detection of ChlaF induced by sunlight across a range of larger scales, from using instruments mounted on towers above plant canopies to Earth-orbiting satellites. This signal is referred to as solar-induced fluorescence (SIF) and its application promises to overcome spatial constraints on studies of photosynthesis, opening new research directions and opportunities in ecology, ecophysiology, biogeochemistry, agriculture and forestry. However, to unleash the full potential of SIF, intensive cross-disciplinary work is required to harmonize these new advances with the rich history of biophysical and ecophysiological studies of ChlaF, fostering the development of next-generation plant physiological and Earth-system models. Here, we introduce the scale-dependent link between SIF and photosynthesis, with an emphasis on seven remaining scientific challenges, and present a roadmap to facilitate future collaborative research towards new applications of SIF.

Leaf density and chemical composition explain variation in leaf mass area with spectral composition among 11 widespread forbs in a common garden.

Leaf mass per area (LMA) is a key leaf functional trait correlated with plant strategies dictating morphology, physiology, and biochemistry. Although sunlight is generally accepted as a dominant factor driving LMA, the contribution of each spectral region of sunlight in shaping LMA is poorly understood. In the present study, we grew 11 widespread forb species in a common garden and dissected the traits underpinning differences in LMA, such as its morphological components (leaf density [LD] and leaf thickness [LT]), macroelement, and metabolite composition under five spectral-attenuation treatments: (1) transmitting c. 95% of the whole solar spectrum (> 280 nm), (2) attenuating ultraviolet-B radiation (UV-B), (3) attenuating both UV-A and UV-B radiation, (4) attenuating UV radiation and blue light, (5) attenuating UV radiation, blue, and green light. We found that LMA, LD, and chemical traits varied significantly across species depending on spectral treatments. LMA was significantly increased by UV-B radiation and green light, while LD was increased by UV-A but decreased by blue light. LMA positively correlated with LD across treatments but was only weakly related to LT, suggesting that LD was a better determinate of LMA for this specific treatment. Regarding leaf elemental and metabolite composition, carbon, nitrogen, and total phenolics were all positively correlated with LMA, whereas lignin, non-structural carbohydrates, and soluble sugars had negative relationships with LMA. These trends imply a tradeoff between biomass allocation to structural and metabolically functional components. In conclusion, sunlight can spectrally drive LMA mainly through modifying functional and structural support.

Alpine forbs rely on different photoprotective strategies during spring snowmelt.

Snowmelt in alpine ecosystems brings ample water, and together with above-freezing temperatures, initiates plant growth. In this scenario, rapid activation of photosynthesis is essential for a successful life-history strategy. But, strong solar radiation in late spring enhances the risk of photodamage, particularly before photosynthesis is fully functional. We compared the photoprotective strategy of five alpine forbs: one geophyte not particularly specialised in subnival life (Crocus albiflorus) and four wintergreens differing in their degree of adaptation to subnival life, from least to most specialised: Gentiana acaulis, Geum montanum, Homogyne alpina and Soldanella alpina. We used distance to the edge of snow patches as a proxy to study time-dependent changes after melting. We postulated that the photoprotective response of snowbed specialists would be stronger than of more-generalist alpine meadow species. Fv /Fm was relatively low across wintergreens and even lower in the geophyte C. albiflorus. This species also had the largest xanthophyll-cycle pool and lowest tocopherol and flavonoid glycoside contents. After snow melting, all the species progressively activated ETR, but particularly the intermediate snowbed species G. acaulis and G. montanum. The photoprotective responses after snowmelt were idiosyncratic: G. montanum rapidly accumulated xanthophyll-cycle pigments, tocopherol and flavonoid glycosides; while S. alpina showed the largest increase in plastochromanol-8 and chlorophyll contents and the greatest changes in optical properties. Climate warming scenarios might shift the snowmelt date and consequently alter the effectiveness of photoprotection mechanisms, potentially changing the fitness outcome of the different strategies adopted by alpine forbs.

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.

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.

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.

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.

Species-specific effect of UV-B radiation on the temporal pattern of leaf growth.

Recent molecular and physiological studies have demonstrated that ultraviolet-B radiation (UV-B) can affect some of the processes involved in leaf growth, but the phases of leaf growth affected have not been clearly delimited. We used functional growth analysis to assess the effects of UV-B radiation on the time course of leaf growth in seedlings of two birch species (Betula pendula and Betula pubescens). Our aim was to identify the phase(s) of leaf development affected by UV-B radiation. In a greenhouse study, 1-year-old birch seedlings were subjected to three daily doses of supplemental UV-B radiation treatments (UV-B⁺) and no UV-B radiation controls (UV-B⁻). Leaf growth measurements every 2 days were complemented by assessment of other functional traits over a 4-week period at the start of the growing season. Using fitted curves, we were able to determine that the rate of leaf expansion was slowed by the UV-B⁺ treatment in leaves of B. pendula because of a slower maximum leaf growth rate compared with plants under the UV-B⁻ controls, but that compensation toward the end of the period of expansion negated this difference when leaves reached their final size. UV-B⁺ had little effect on the rate of B. pubescens leaf growth despite a larger reduction in leaf final size due to UV-B⁺ than occurred in B. pendula leaves. In conclusion, effective regulation ameliorated the effects of UV-B radiation on leaf and seedling growth in B. pendula, whereas in B. pubescens, reductions in leaf final size under UV-B⁺ were consistent with a slightly reduced rate of height growth.