Prolonged exposure to hypergravity increases number and size of cells and enhances lignin deposition in the stem of Arabidopsis thaliana.

We have performed a lab-based hypergravity cultivation experiment using a centrifuge equipped with a lighting system and examined long-term effects of hypergravity on the development of the main axis of the Arabidopsis (Arabidopsis thaliana (L.) Heynh.) primary inflorescence, which comprises the rachis and peduncle, collectively referred to as the main stem for simplicity. Plants grown under 1 × g (gravitational acceleration on Earth) conditions for 20-23 days and having the first visible flower bud were exposed to hypergravity at 8 × g for 10 days. We analyzed the effect of prolonged hypergravity conditions on growth, lignin deposition, and tissue anatomy of the main stem. As a result, the length of the main stem decreased and cross-sectional area, dry mass per unit length, cell number, and lignin content of the main stem significantly increased under hypergravity. Lignin content in the rosette leaves also increased when they were exposed to hypergravity during their development. Except for interfascicular fibers, cross-sectional areas of the tissues composing the internode significantly increased under hypergravity in most types of the tissues in the basal part than the apical part of the main stem, indicating that the effect of hypergravity is more pronounced in the basal part than the apical part. The number of cells in the fascicular cambium and xylem significantly increased under hypergravity both in the apical and basal internodes of the main stem, indicating a possibility that hypergravity stimulates procambium activity to produce xylem element more than phloem element. The main stem was suggested to be strengthened through changes in its morphological characteristics as well as lignin deposition under prolonged hypergravity conditions.

Leaf optical properties and photosynthesis of fern species with a wide range of divergence time in relation to mesophyll anatomy.

BACKGROUND AND AIMS: For a comprehensive understanding of the mechanisms of changing plant photosynthetic capacity during plant evolutionary history, knowledge of leaf gas exchange and optical properties are essential, both of which relate strongly to mesophyll anatomy. Although ferns are suitable for investigating the evolutionary history of photosynthetic capacity, comprehensive research of fern species has yet to be undertaken in this regard. METHODS: We investigated leaf optical properties, gas exchange and mesophyll anatomy of fern species with a wide range of divergence time, using 66 ferns from natural habitats and eight glasshouse-grown ferns. We used a spectroradiometer and an integrating sphere to measure light absorptance and reflectance by the leaves. KEY RESULTS: The more newly divergent fern species had a thicker mesophyll, a larger surface area of chloroplasts facing the intercellular airspaces (Sc), thicker cell walls and large light absorptance. Although no trend with divergence time was obtained in leaf photosynthetic capacity on a leaf-area basis, when the traits were expressed on a mesophyll-thickness basis, trends in leaf photosynthetic capacity became apparent. On a mesophyll-thickness basis, the more newly divergent species had a low maximum photosynthesis rate, accompanied by a low Sc. CONCLUSIONS: We found a strong link between light capture, mesophyll anatomy and photosynthesis rate in fern species for the first time. The thick mesophyll of the more newly divergent ferns does not necessarily relate to the high photosynthetic capacity on a leaf-area basis. Rather, the thick mesophyll accompanied by thick cell walls allowed the ferns to adapt to a wider range of environments through increasing leaf toughness, which would contribute to the diversification of fern species.

How plants grow under gravity conditions besides 1 g: perspectives from hypergravity and space experiments that employ bryophytes as a model organism.

Plants have evolved and grown under the selection pressure of gravitational force at 1 g on Earth. In response to this selection pressure, plants have acquired gravitropism to sense gravity and change their growth direction. In addition, plants also adjust their morphogenesis in response to different gravitational forces in a phenomenon known as gravity resistance. However, the gravity resistance phenomenon in plants is poorly understood due to the prevalence of 1 g gravitational force on Earth: not only it is difficult to culture plants at gravity > 1 g(hypergravity) for a long period of time but it is also impossible to create a < 1 genvironment (µg, micro g) on Earth without specialized facilities. Despite these technical challenges, it is important to understand how plants grow in different gravity conditions in order to understand land plant adaptation to the 1 g environment or for outer space exploration. To address this, we have developed a centrifugal device for a prolonged duration of plant culture in hypergravity conditions, and a project to grow plants under the µg environment in the International Space Station is also underway. Our plant material of choice is Physcomitrium (Physcomitrella) patens, one of the pioneer plants on land and a model bryophyte often used in plant biology. In this review, we summarize our latest findings regarding P. patens growth response to hypergravity, with reference to our on-going "Space moss" project. In our ground-based hypergravity experiments, we analyzed the morphological and physiological changes and found unexpected increments of chloroplast size and photosynthesis rate, which might underlie the enhancement of growth and increase in the number of gametophores and rhizoids. We further discussed our approaches at the cellular level and compare the gravity resistance in mosses and that in angiosperms. Finally, we highlight the advantages and perspectives from the space experiments and conclude that research with bryophytes is beneficial to comprehensively and precisely understand gravitational responses in plants.

Correction to: Why is chlorophyll b only used in light-harvesting systems?

The article Why is chlorophyll b only used in light-harvesting systems.

Correction to: Importance of the green color, absorption gradient, and spectral absorption of chloroplasts for the radiative energy balance of leaves.

The article "Importance of the green color, absorption gradient, and spectral absorption of chloroplasts for the radiative energy balance of leaves", written by Atsushi Kume, was originally published Online First without open access.

Why is chlorophyll b only used in light-harvesting systems?

Chlorophylls (Chl) are important pigments in plants that are used to absorb photons and release electrons. There are several types of Chls but terrestrial plants only possess two of these: Chls a and b. The two pigments form light-harvesting Chl a/b-binding protein complexes (LHC), which absorb most of the light. The peak wavelengths of the absorption spectra of Chls a and b differ by c. 20 nm, and the ratio between them (the a/b ratio) is an important determinant of the light absorption efficiency of photosynthesis (i.e., the antenna size). Here, we investigated why Chl b is used in LHCs rather than other light-absorbing pigments that can be used for photosynthesis by considering the solar radiation spectrum under field conditions. We found that direct and diffuse solar radiation (PARdir and PARdiff, respectively) have different spectral distributions, showing maximum spectral photon flux densities (SPFD) at c. 680 and 460 nm, respectively, during the daytime. The spectral absorbance spectra of Chls a and b functioned complementary to each other, and the absorbance peaks of Chl b were nested within those of Chl a. The absorption peak in the short wavelength region of Chl b in the proteinaceous environment occurred at c. 460 nm, making it suitable for absorbing the PARdiff, but not suitable for avoiding the high spectral irradiance (SIR) waveband of PARdir. In contrast, Chl a effectively avoided the high SPFD and/or high SIR waveband. The absorption spectra of photosynthetic complexes were negatively correlated with SPFD spectra, but LHCs with low a/b ratios were more positively correlated with SIR spectra. These findings indicate that the spectra of the photosynthetic pigments and constructed photosystems and antenna proteins significantly align with the terrestrial solar spectra to allow the safe and efficient use of solar radiation.

Nutrient deficiency promotes male-biased apparent sex ratios at the ramet level in the dioecious plant Myrica gale var. tomentosa in oligotrophic environments in bogs.

In populations of dioecious plants, the differences in the cost of reproduction between male and female plants can promote a male-biased sex ratio. In this study, we examine the macronutrient levels in tissues of the dioecious wetland shrub Myrica gale to identify the cost of reproduction for male and female plants and to examine the effect of nutrients on the apparent sex ratio at the ramet level. We examined plants across 12 populations of M. gale inhabiting bogs and fens in Japan. For each population, we used line transects to estimate the apparent sex ratio and measured the concentrations of nitrogen (N), phosphorus (P), and potassium (K) in the leaves sampled from male and female plants and in the fruits from female plants. For five of the populations, we calculated the flowering frequency, mortality, and the recruitment rate (as the rate of clonal propagation). We found that the proportion of females was positively affected, and the male bias of sex ratios reduced, by increases in P concentration in leaves sampled from female plants. Neither mortality nor recruitment was affected by sex or by the nutrient concentration (P, K). The flowering frequency was not affected by sex or by K concentration, but decreased with decreases in the P concentration measured in leaves. This study confirmed that reproduction in M. gale is P-limited. We found no distinct differences in the flowering frequency, mortality, or recruitment rate between the male and female plants.

Correction to: A hypergravity environment increases chloroplast size, photosynthesis, and plant growth in the moss Physcomitrella patens.

The original article can be found online.

Importance of the green color, absorption gradient, and spectral absorption of chloroplasts for the radiative energy balance of leaves.

Terrestrial green plants absorb photosynthetically active radiation (PAR; 400-700 nm) but do not absorb photons evenly across the PAR waveband. The spectral absorbance of photosystems and chloroplasts is lowest for green light, which occurs within the highest irradiance waveband of direct solar radiation. We demonstrate a close relationship between this phenomenon and the safe and efficient utilization of direct solar radiation in simple biophysiological models. The effects of spectral absorptance on the photon and irradiance absorption processes are evaluated using the spectra of direct and diffuse solar radiation. The radiation absorption of a leaf arises as a consequence of the absorption of chloroplasts. The photon absorption of chloroplasts is strongly dependent on the distribution of pigment concentrations and their absorbance spectra. While chloroplast movements in response to light are important mechanisms controlling PAR absorption, they are not effective for green light because chloroplasts have the lowest spectral absorptance in the waveband. With the development of palisade tissue, the incident photons per total palisade cell surface area and the absorbed photons per chloroplast decrease. The spectral absorbance of carotenoids is effective in eliminating shortwave PAR (<520 nm), which contains much of the surplus energy that is not used for photosynthesis and is dissipated as heat. The PAR absorptance of a whole leaf shows no substantial difference based on the spectra of direct or diffuse solar radiation. However, most of the near infrared radiation is unabsorbed and heat stress is greatly reduced. The incident solar radiation is too strong to be utilized for photosynthesis under the current CO2 concentration in the terrestrial environment. Therefore, the photon absorption of a whole leaf is efficiently regulated by photosynthetic pigments with low spectral absorptance in the highest irradiance waveband and through a combination of pigment density distribution and leaf anatomical structures.

A hypergravity environment increases chloroplast size, photosynthesis, and plant growth in the moss Physcomitrella patens.

The physiological and anatomical responses of bryophytes to altered gravity conditions will provide crucial information for estimating how plant physiological traits have evolved to adapt to significant increases in the effects of gravity in land plant history. We quantified changes in plant growth and photosynthesis in the model plant of mosses, Physcomitrella patens, grown under a hypergravity environment for 25 days or 8 weeks using a custom-built centrifuge equipped with a lighting system. This is the first study to examine the response of bryophytes to hypergravity conditions. Canopy-based plant growth was significantly increased at 10×g, and was strongly affected by increases in plant numbers. Rhizoid lengths for individual gametophores were significantly increased at 10×g. Chloroplast diameters (major axis) and thicknesses (minor axis) in the leaves of P. patens were also increased at 10×g. The area-based photosynthesis rate of P. patens was also enhanced at 10×g. Increases in shoot numbers and chloroplast sizes may elevate the area-based photosynthesis rate under hypergravity conditions. We observed a decrease in leaf cell wall thickness under hypergravity conditions, which is in contrast to previous findings obtained using angiosperms. Since mosses including P. patens live in dense populations, an increase in canopy-based plant numbers may be effective to enhance the toughness of the population, and, thus, represents an effective adaptation strategy to a hypergravity environment for P. patens.

Leaf color is fine-tuned on the solar spectra to avoid strand direct solar radiation.

The spectral distributions of light absorption rates by intact leaves are notably different from the incident solar radiation spectra, for reasons that remain elusive. Incident global radiation comprises two main components; direct radiation from the direction of the sun, and diffuse radiation, which is sunlight scattered by molecules, aerosols and clouds. Both irradiance and photon flux density spectra differ between direct and diffuse radiation in their magnitude and profile. However, most research has assumed that the spectra of photosynthetically active radiation (PAR) can be averaged, without considering the radiation classes. We used paired spectroradiometers to sample direct and diffuse solar radiation, and obtained relationships between the PAR spectra and the absorption spectra of photosynthetic pigments and organs. As monomers in solvent, the spectral absorbance of Chl a decreased with the increased spectral irradiance (W m(-2) nm(-1)) of global PAR at noon (R(2) = 0.76), and was suitable to avoid strong spectral irradiance (λmax = 480 nm) rather than absorb photon flux density (µmol m(-2) s(-1) nm(-1)) efficiently. The spectral absorption of photosystems and the intact thallus and leaves decreased linearly with the increased spectral irradiance of direct PAR at noon (I dir-max), where the wavelength was within the 450-650 nm range (R(2) = 0.81). The higher-order structure of photosystems systematically avoided the strong spectral irradiance of I dir-max. However, when whole leaves were considered, leaf anatomical structure and light scattering in leaf tissues made the leaves grey bodies for PAR and enabled high PAR use efficiency. Terrestrial green plants are fine-tuned to spectral dynamics of incident solar radiation and PAR absorption is increased in various structural hierarchies.

Correction: Evaluation of thermoregulation of different pine organs in early spring and estimation of heat reward for the western conifer seed bug (Leptoglossus occidentalis) on male cones.

[This corrects the article DOI: 10.1371/journal.pone.0272565.].

Life Cycle Plasticity in Typhula and Pistillaria in the Arctic and the Temperate Zone.

Typhulaceae Jülich is one of the cold-adapted fungal families in basidiomycetes. The representative genera, Typhula (Pers.) Fr. and Pistillaria Fr., are distinguished by the discontinuity between stems and hymenia in the former and the continuity in the latter (Fries 1821). This taxonomic criterion is ambiguous, and consequently, the view of Karsten (1882) has been widely accepted: Typhula develops basidiomata from sclerotia, while basidiomata develop directly from substrata in Pistillaris. However, Corner (1970) observed basidiomata of Pistillaria petasitis S. Imai developing from sclerotia in Hokkaido, Japan. We later recognized that P. petasitis basidiomata also emerged directly from substrates on the ground in Hokkaido. An aberrant form of Typhula hyperborea H. Ekstr. was found in Upernavik, West Greenland. This specimen had a stem-like structure on a Poaceae plant, and sclerotia developed on its tip. Similar phenomena were found in other Typhula species in Japan. In this study, we aimed to elucidate the life cycle plasticity in the genera Typhula and Pistillaria through the interactions between their ecophysiological potential and environmental conditions in their localities. We collected and prepared strains of the above fungi from sclerotia or basidiomata, and we elucidated the taxonomical relationship and determined the physiological characteristics of our strains. Our findings imply that both Typhula and Pistillaria have the potential to produce sclerotia as well as the capacity for mycelial growth at ambient air temperatures in each locality where samples were collected. These findings suggest that Typhula spp. develope basidiomata not only from the sclerotia dispersed by the basidiospores but also from mycelia generated by the spore germination, which formed basidiomata multiple times, depending on their growth environments.

Nanodiamond-Enhanced Nanofiber Separators for High-Energy Lithium-Ion Batteries.

Current lithium-ion battery separators made from polyolefins such as polypropylene and polyethylene generally suffer from low porosity, low wettability, and slow ionic conductivity and tend to perform poorly against heat-triggering reactions that may cause potentially catastrophic issues, such as fire. To overcome these limitations, here we report that a porous composite membrane consisting of poly(vinylidene fluoride-co-hexafluoropropylene) nanofibers functionalized with nanodiamonds (NDs) can realize a thermally resistant, mechanically robust, and ionically conductive separator. We critically reveal the role of NDs in the polymer matrix of the membrane to improve the thermal, mechanical, crystalline, and electrochemical properties of the composites. Taking advantages of these characteristics, the ND-functionalized nanofiber separator enables high-capacity and stable cycling of lithium cells with LiNi0.8Mn0.1Co0.1O2 (NMC811) as the cathode, much superior to those using conventional polyolefin separators in otherwise identical cells.

Impact of soil water chemistry on the apparent sex ratio of the flowering ramets of the dioecious plant Myrica gale var. tomentosa.

We determined whether the apparent (M/Fl) sex ratio (male ramets/flowering ramets) and apparent reproductive ramet ratio (Fl/Li ratio; flowering ramets/living ramets) in 15 Myrica gale var. tomentosa populations varied with dissolved total nitrogen, dissolved total phosphorus, potassium, magnesium, calcium, or pH in the soil water. Our aim was to define the environmental factors affecting the M/Fl sex ratio and Fl/Li ratio of the populations. We also examined the habitat conditions of these populations by analyzing soil water chemistry and water dynamics. In 2007, 3 of the 15 populations had no females. The remaining 12 had significantly male-biased (M/Fl sex ratio = 0.59-0.97). Although we could not explain the absence of females by the current potassium levels alone, as potassium increased, so did the M/Fl sex ratio. As nitrogen increased and potassium decreased, Fl/Li ratio decreased. Our soil water chemistry analyses suggested that the potassium supply by soil surface erosion from flooding and the inflow of anthropogenic nitrogen were the important factors influencing the M/Fl sex ratio and Fl/Li ratio. Nitrogen management would be important in one of the endangered populations where inflow of nitrogen was the highest among 15 habitats.

Effects of sex and soil water chemistry on leaf morphology and physiology of Myrica gale var. tomentosa.

Plants respond to environmental stressors, such as an oligotrophic environments, by altering the morphological and physiological functions of their leaves. Sex affects these functions because of the asymmetric cost of reproduction in dioecious plants. We compared the leaf mass per leaf area (LMA), ratio of intercellular air space in leaf mesophyll tissue (mesophyll porosity), palisade thickness, and carbon isotope ratio (δ13C) of leaves of the dioecious shrub Myrica gale based on sex and gradients of soil water chemistry across habitats in the field. The PCA showed that the first three principal components accounted for 84.5% of the variation. PC1 to PC3 were associated with the origin of soil water, nitrogen status of habitats, and sea-salt contributions, respectively. LMA varied from 5.22 to 7.13 µg/cm2, and it was positively related to PC2 and negatively related to PC3, but not to PC1 or sex, suggesting that LMA was low under poor nitrogen conditions and varied with salinity. Mesophyll porosity values were over 50% for all habitats. Mesophyll porosity was positively affected by PC3 and smaller in females than in males. This suggests that M. gale exhibits differences in mesophyll anatomy according to sex. Palisade thickness ranged from 0.466 to 0.559 mm/mm. The leaves of females had thinner palisade layers per mesophyll layer than those of males; however, the habitat did not affect the thickness of the palisade layer per mesophyll layer. The δ13C values of leaves varied from -32.14 to -30.51 ‰. We found that δ13C values were positively related to PC2 but not to PC1, PC3, and sex. Under poor nitrogen conditions, the δ13C of M. gale leaves decreased, suggesting that nutrient deficiency would decrease more under the long-term averaged ratio of photosynthesis than stomatal conductance, leading to low water use efficiency.

Evaluation of thermoregulation of different pine organs in early spring and estimation of heat reward for the western conifer seed bug (Leptoglossus occidentalis) on male cones.

The western conifer seed bug (WCSB, Leptoglossus occidentalis) is a pest of many pine species and is invasive worldwide. WCSB directly and indirectly deteriorates pine nut production by sucking seeds from cones. Currently, researchers think that WCSBs search for food by a combination of cues from visible light, infrared radiation, and chemicals such as monoterpenes. Some research revealed that WCSBs prefer larger cones, and it was thought that WCSBs suck seeds from and obtain more heat on larger cones. However, in early spring, we observed that most WCSBs gathered on male cones rather than on female cones and young cones. We hypothesized that male pine cones were warmer than female cones and needles, and WCSBs sucking male cones may receive more heat. To test these hypotheses, we measured spectral reflectance with a hyperspectral sensor and temperature of pine organs with tiny thermocouples, and the data were analyzed by a heat budget model. Our results revealed that male cones were significantly warmer and more reflective than female cones and needles, which may attract WCSBs. These results supported our hypothesis that WCSBs on male cones were warmer than those on other organs. This study will help further understanding of WCSBs and the adaptive value of pine cone colors.

Three-dimensionally visualized rhizoid system of moss, Physcomitrium patens, by refraction-contrast X-ray micro-computed tomography.

Land plants have two types of shoot-supporting systems, root system and rhizoid system, in vascular plants and bryophytes. However, since the evolutionary origin of the systems is different, how much they exploit common systems or distinct systems to architect their structures is largely unknown. To understand the regulatory mechanism of how bryophytes architect the rhizoid system responding to environmental factors, we have developed the methodology to visualize and quantitatively analyze the rhizoid system of the moss, Physcomitrium patens, in 3D. The rhizoids having a diameter of 21.3 µm on the average were visualized by refraction-contrast X-ray micro-computed tomography using coherent X-ray optics available at synchrotron radiation facility SPring-8. Three types of shape (ring-shape, line and black circle) observed in tomographic slices of specimens embedded in paraffin were confirmed to be the rhizoids by optical and electron microscopy. Comprehensive automatic segmentation of the rhizoids, which appeared in three different form types in tomograms, was tested by a method using a Canny edge detector or machine learning. The accuracy of output images was evaluated by comparing with the manually segmented ground truth images using measures such as F1 score and Intersection over Union, revealing that the automatic segmentation using machine learning was more effective than that using the Canny edge detector. Thus, machine learning-based skeletonized 3D model revealed quite dense distribution of rhizoids. We successfully visualized the moss rhizoid system in 3D for the first time.

The ratio of transmitted near-infrared radiation to photosynthetically active radiation (PAR) increases in proportion to the adsorbed PAR in the canopy.

The daily total photosynthetically active radiation (400-700 nm, PAR) and near-infrared radiation (700-1000 nm, NIR) were measured in the understory beneath the canopy (PARt and NIRt) and above the canopy (PARi and NIRi) of a Japanese cool-temperate deciduous broad-leaved forest during the snow-free period (May to November). The integration of spectral radiation for NIR and that for PAR, and the daily integrations of instantaneous NIR and PAR, reduced the noises from the optical difference in spectrum and from canopy structure heterogeneity, sky condition and solar elevation. PARi/PARt was linearly related to NIRt/PARt (R² = 0.96). The effect of cloudiness was negligible, because the fluctuation of NIRi/PARi was quite small regardless of season and weather conditions compared with the range of NIRt/PARt in the forest. The ratio of NIRt/PARt beneath the canopy was log-linearly related to the in situ leaf area index (LAI) with a wide range from 0 to 5.25 (R² = 0.97). We conclude that seasonal changes in fAPAR (= 1 - PARt/PARi) and LAI of a canopy can be estimated with high accuracy by transmitted NIRt and PARt beneath the canopy.

Number concentration and size distribution of ultrafine particles on the roadside of the Tateyama-Kurobe Alpine route, Japan.

Measurements of ultrafine particles (UFPs) with diameters smaller than 100 nm were made with a Scanning Mobility Particle Sizer on the roadside of the Tateyama-Kurobe Alpine route on the western slope of Mt. Tateyama, Japan, in the summer and autumn of 2007 and 2008. The number concentrations of nanoparticles (NPs) with diameters smaller than 50 nm increased sharply in the morning. This increase agreed with that of the NO concentration, which is a good indicator of vehicle emissions. Although the peak concentration of NO was also detected in the late afternoon, the concentrations of NPs did not increase. The vehicle emission of UFPs may vary considerably with the driving conditions. The emission of NPs might have been accelerated under a high engine-load condition. The NP concentrations in October were much higher than those in August, although there were more buses in August than in October. The number size distribution in the morning was bimodal, with the first peak around 20-30 nm and the second one around 70-80 nm. The first peak became small during the daytime. The first peak disappeared, and a small second peak remained through midnight.