Lycophytes, one of the earliest-diverging vascular plant lineages, occur in diverse and often extreme environments, yet their ecological adaptation strategies remain poorly understood. Ionomics, characterized by elemental composition, offers insights into how evolutionary history and environmental conditions shape plant function. Here, we investigated how lineage and soil type influence ionomic profiles in terrestrial lycophytes by analyzing shoot elemental concentrations in three Lycopodiaceae species and four Selaginellaceae species collected across Japan, including serpentine soils and solfatara fields. Element concentrations were quantified by inductively coupled plasma mass spectrometry, with nitrogen and phosphorus measured colorimetrically. Statistical analyses included heatmap clustering and principal component analysis. Across non-solfatara environments, Lycopodiaceae and Selaginellaceae showed distinct ionomic profiles at the family level within the sampled taxa, with variation appearing to be more strongly affected by lineage differences than by living environment. In Selaginellaceae, individuals from serpentine soils showed higher magnesium, iron, and arsenic, reflecting serpentine soil characteristics. Lycopodiaceae showed higher aluminum, copper, potassium, rubidium, and manganese, whereas Selaginellaceae had higher calcium, magnesium, phosphorus, nitrogen, strontium, and zinc. For Palhinhaea cernua (Lycopodiaceae), which occurred in both solfatara and non-solfatara fields, shoots from the acidic solfatara environment contained higher sulfur, nitrogen, and phosphorus while maintaining consistently high aluminum concentrations, supporting its status as an aluminum hyperaccumulator. Overall, lycophytes demonstrate family-specific elemental homeostasis that is conserved across environments, alongside characteristic shifts in extreme soils. These findings provide a foundation for understanding ecological diversification in the earliest-diverging vascular plants and highlight ionomics as an effective approach for exploring their environmental adaptation strategies.
Methane (CH4) is the second most important greenhouse gas after carbon dioxide (CO2), and its atmospheric concentration is on the rise. Soil CH4 consumption (=absorption) capacity is declining due to reduced forests and green spaces, as well as other environmental factors and anaerobic stresses. Environmental and stand structure parameters were cross-referenced with publicly available international ecosystem databases, such as FLUXNET, ICOS, NEON, AmeriFlux, the TRY plant trait database and the Oak Ridge FACE site. Searches were conducted using keywords such as region, water level, and stand density. The data indicate that under high-CO2 conditions, the increase of forest canopy density leads to increased litter accumulation on the forest floor and reduced sunlight penetration, creating anaerobic conditions. This can cause forests to shift from CH4 consumption to CH4 release. Based on these findings, we discussed methods to maintain and enhance the CH4-absorbing capacity of forest soils. This can be achieved through management practices that improve environmental conditions and increase soil fauna’s activity, such as those associated with thinning operations in overmature forest stands across various regions. This ecological manipulation through thinning practices promotes ground-level temperature increases and the activities of soil fauna, as well as maintaining aerobic conditions near the soil surface.
ABSTRACT Acidic soils impose multiple stresses on plants and affect soil nutrient availability. Some elements are essential to many physiological processes within plants. Comprehensive studies on the ionomic profiles in plants are insightful for uncovering plant strategies for nutrient uptake and tolerance to environmental stress. This study aimed to elucidate plant nutrition strategies by comparing the ionomes of plants grown in highly acidic soils, so‐called solfatara fields, and in non‐solfatara fields, with average pH values of 3.89 and 5.29, respectively. A quadrat survey was conducted to investigate the effects of fumaroles, which emit volcanic gases and are characteristic of solfatara fields, on plant ionomes. Our study classified ionomic profiles in solfatara and non‐solfatara fields into two groups: aluminum (Al)‐accumulating and non‐Al‐accumulating plants. Among non‐Al‐accumulating plants, concentrations of manganese, magnesium, and sulfur were similar or higher in solfatara than in non‐solfatara fields. Among Al‐accumulating plants, the concentration of Al in Palhinhaea cernua was higher than in other species, and it did not differ significantly between solfatara and non‐solfatara fields. In the quadrat survey, only four species were found, and their ionomes did not change with distance from the fumarole. The most acidic area had a pH of 2.28, and only Fimbristylis dichotoma subsp. podocarpa was found there. Although it remains unclear whether these patterns reflect species traits or adaptive responses to environmental conditions, the results enable us to hypothesize that manganese, magnesium, and sulfur were associated with tolerance for very acidic soil exhibited by non‐Al‐accumulating plants.
Manganese (Mn) hyperaccumulator plants are rare in nature but provide important insights into plant-soil interactions, Mn nutritional physiology, and biogeochemical cycling. X-ray fluorescence (XRF) herbarium analysis of Japanese wild plants was used to investigate Mn concentrations in 25 different Ilex species (totaling 619 specimens). Among the 25 analyzed Ilex species, nine species had leaves containing > 10,000 mg kg(-1) Mn (dry weight), and in six of these species, multiple hyperaccumulator-level specimens were found, representing 8%-21% of the specimens within those six species. Analysis of Mn and six other essential elements (K, Ca, P, S, Fe, and Zn) revealed that Ilex had, on average, four times higher Zn concentrations than those in most other plants, with a significant positive correlation between Mn and Zn. No other elements showed significant correlations with Mn. In contrast to other Mn hyperaccumulators, including Chengiopanax sciadophylloides from Japan, no positive Mn-Ca correlation was observed. These findings identify Ilex as the second Mn hyperaccumulator group reported from Japan and show that, like C. sciadophylloides, hyperaccumulation in Ilex was achieved under ordinary forest soils without known contamination sources (noncontaminated soil). The distinct accumulation pattern observed here in the genus Ilex suggests a physiological mechanism different from the Mn-Ca co-accumulation observed in other Mn hyperaccumulators.
For the use of phosphorus (P) in slag, a steel production by-product, as an agricultural fertilizer, we conducted pot cultivation experiments using prototype P concentrated slags as a P source. Maize (Zea mays L.) plants were cultivated as a main crop for 32 days to elucidate mineral uptake by crop plants in a pot culture system. Effects of white lupin (Lupinus albus L.) and blue lupin (L. angustifolius L.) intercropping on mineral uptake by maize were also examined. Growth and P content of monocropped maize were high in the KHP-02 treatment, which used slag with high P availability, and were comparable with those achieved using fused phosphate. However, P uptake by maize was low in the SHP27 treatment, with its lower P availability slag. Growth and P content in maize with applied KHP-02 and fused phosphate were significantly high in intercropping with blue lupin. The pH of post-cultivation pot soil was significantly lower in both lupin species than in maize. These findings suggest that P in the rhizosphere was solubilized by lupin root exudates. The P fractionation of whole soil after cultivation revealed that soluble fractions in the intercropped pots were lower than in monocropped pots of maize, suggesting that the mobilized P in the rhizosphere was accumulated immediately by intercropped plants. Multi-element analysis revealed higher accumulation not only of P but also of other elements such as calcium, silicon, magnesium, iron, and molybdenum than in treatments of KHP-02 and fused phosphate. Ion-profile-based principal component analysis showed that the fertilizer had a strong effect, but no clear differences were found in maize among planting system differences. In conclusion, the P concentrated slag fertilizer has the potential to provide P for crops at levels similar to those obtained using conventional fertilizers. Findings suggest that organic acids in the root exudates of lupins can improve P absorption from slag-derived P.
Enhanced Rock Weathering (ERW) is a negative emission technology in which crushed rock powder is applied to cropland to sequestrate carbon by enhancing chemical weathering of silicate minerals. It is a co-benefit approach to promote crop growth by supplying elements (silicon (Si), calcium (Ca), magnesium (Mg), etc.) from weathering rock as well. In this study, we cultivated paddy rice for 99 days in a pot experiment with Gray Lowland soil (pH = 6.19) with six application rates of basalt equivalent to 0, 5, 10, 20, 50, 100 t ha(-1). We investigated the change of soil chemical properties and elemental uptake in rice. Then, we calculated the release of Si, Ca, Mg, potassium (K), and sodium (Na) from the basalt, and carbon balance between before planting and after harvest in the whole system including soil and plant. The application of the basalt increased the total amount of soil available Si and exchangeable Mg and Na in the pot as well as the uptake of Si and Na in rice (p < 0.05). There was no significant difference in arsenic, cadmium, chromium, copper, and nickel concentrations in rice grain among the application rates of the basalt. The release of Si, Mg, and Na from the basalt increased corresponding to the application rate of the basalt, suggesting that the applied basalt was weathered during cultivation, though increased carbon sequestration by the basalt weathering was not detected in 99 days of this experiment. We clearly demonstrated that basalt powder got weathered and released Si, Mg, and Na in paddy field condition and additional Si enhanced paddy rice growth.
Paperpots have been introduced as biodegradable containers for agricultural seedlings. Nevertheless, to prevent the premature decomposition of paper by soil microorganisms during seedling growth, these pots are often reinforced with synthetic fibers, chemically modified, or treated with fungicides. However, these interventions result in the retention of non-biodegradable residues in the field post-transplantation. To address this, we studied the controlled decay of decay-resistant citric acid cross-linked paper and evaluated its use for making paperpots for agricultural use. Decay-resistant paper was prepared using citric acid and sodium hypophosphite, followed by heating and alkali treatments. For the control, dimethylol dihydroxyethylene urea (DMDHEU) -treated paper was similarly prepared. The paper's wet tensile strength was assessed after exposure to water, cellulase, and soil biodegradation. Additionally, paperpots were manufactured and used to grow Komatsuna seedlings, with and without alkali treatment, to evaluate the control of biodegradability and the impact on seedling growth. Treating the decay-resistant citric acid cross-linked paper with alkali, such as calcium hydroxide, which is used as often used in agriculture, cleaved the ester bonds formed by the hydroxyl groups of cellulose and citric acid, resulting in a paper that could decompose. Treating Komatsuna seedlings grown in citric acid cross-linked paperpots with alkali treatment resulted in pot decomposition without affecting the growth of Komatsuna. Our results showed that controlled decay of citric acid cross-linked paperpots can be achieved by treating them with an alkaline agent like calcium hydroxide, which is routinely used in agriculture, without affecting plant growth.
Climate change affects food production, increasing the need for CO2 removal (CDR) strategies. Enhanced rock weathering, which involves the spreading of crushed silicate rock powder on agricultural soil to sequester atmospheric CO2 via enhanced natural weathering, shows potential to enrich the agricultural soil. In this study, we evaluated the short-term impacts of basalt powder on CO2 emissions in rhizosphere and non-rhizosphere soils and estimated the field carbon budget in an experimental soybean field at Hokkaido University, Japan. Basalt powder application at 150 Mg ha−1, with incorporation into the soil to a depth of 15 cm, significantly increased the soil pH and reduced the soil volumetric water content. Regardless of treatment, the carbon budget was negative, indicating the overall carbon loss in field. Basalt powder application reduced this carbon loss from 2.69 ± 0.41 to 1.90 ± 0.73 Mg C ha−1, primarily by absorbing CO2 that would otherwise have been released into the atmosphere through weathering and sequestering it in the soil, though the difference was not significant. ERW-induced CO2 emission reduction rate was 0.81 ± 0.17 Mg C ha−1, with approximately 86.4
Hakea laurina, a woody Proteaceae, naturally occurs in severely phosphorus (P)-impoverished habitats in southwest Australia. It develops distinctive cluster roots that exhibit a high capacity for carboxylate exudation and acid phosphatase activity, contributing to its P acquisition. However, the molecular mechanisms underlying these physiological functions remain poorly understood. We explored the cluster-root transcriptome using de novo RNA-Seq and identified Hakea laurina Aluminum-activated Malate Transporter 1 (HalALMT1), encoding an aluminum (Al)-activated malate transporter induced in mature cluster roots. We characterized HalALMT1 through electrophysiological assays and overexpression in Arabidopsis thaliana, and localized HalALMT1 expression, acid phosphatase activity, and suberized boundaries in cluster roots. Differentially expressed genes highlighted multiple increased carboxylate-related processes at cluster-root maturity. HalALMT1 released malate, an activity further enhanced by exposure to Al3+. Notably, HalALMT1 was specifically expressed in mature cortex cells of cluster rootlets, which lack a suberized exodermis. Acid phosphatase activity was pronounced throughout the cluster rootlets, unlike in noncluster roots where it was limited to the epidermis and stele. Substantial malate release and acid phosphatase activity in the cortex cells in cluster rootlets, which lack a suberized exodermis, allowed massive exudation. This study sheds light on an exquisite P-acquisition strategy of Proteaceae, enabling survival under extremely low P availability.
The growth of Larix species has been improved by hybridization, while the hybrid with salt-intolerant species may not be useful for afforestation in saline soil. This study evaluated the relationship between growth responses to salt stress and root functional traits in seedlings of L. gmelinii var. japonica, L. kaempferi, and their hybrid (L. gmelinii var. japonica × L. kaempferi). In a greenhouse, two-year-old seedlings were cultivated with 70 mM of NaCl loading. Roots were divided into tap roots, lignified woody lateral roots, and nonlignified feeder lateral roots. The dry mass, total length and total area and Na concentration of each root were evaluated. The total dry mass of hybrid larch was higher than L. gmelinii regardless of salt stress. The total dry mass and total length of a tap root was suppressed in L. kaempferi by NaCl loading, but not in the other two species. These results demonstrated that the variation in salt tolerance was associated with the vertical distribution of root growth. While Na concentration in primary roots was increased in L. kaempferi and hybrid larch under NaCl loading, the dry mass of primary roots was reduced only in L. kaempferi. In L. gmelinii, dry mass in a tap root and secondary roots were not decreased with the highest Na concentration, indicating the tolerance to Na toxicity accumulated in these roots. Hybrid larch showed no growth suppression with increasing Na concentration in primary roots, suggesting that its superior salt tolerance would be inherited from the mother parent.
Paprika (Capsicum annuum L.) is known for its abundant functional components such as carotenoids and phenolics. Growth responses to nutrient conditions including nitrogen (N) have been studied. However, there has been only limited analysis on the variations in components in response to N. This study investigated the effects of different N levels on the functional components of paprika. Paprika was grown under different N conditions and ripe fruits were harvested. Subsequently, mineral, carotenoid, phenolics, and sugar contents were measured. Higher N supply increased fruit weight and carotenoids, but reduced phenolics and glucose. The negative correlation between carotenoids and phenolics suggests complementary antioxidant functions in fruits, while the significant association between these metabolites and boron and potassium implies that these elements may have indirectly influenced metabolic pathways. Moreover, specific components like phenolics indicate a trade-off with yield. Capsanthin alone was significantly affected by N supply, suggesting downstream effects on the carotenoid biosynthetic pathway. This study will contribute to the optimization of N fertilizer application for balanced paprika production with high quality and economic yield.
Recent studies suggest that biochar may enhance biological nitrogen fixation (BNF) in legumes, yet the underlying mechanisms remain unclear. This study aimed to explore the potential effects of manure-derived biochar on BNF, with a focus on its influence on soil physicochemical properties and the availability of essential nutrients such as molybdenum (Mo), a cofactor for nitrogenase. A greenhouse experiment was conducted using dairy manure biochar or lime applied to soils growing white clover (Trifolium repens), either with or without phosphorus (P) and potassium (K) fertilizers. Plant growth, nutrient uptake, nitrogenase activity, and post-harvest soil properties were evaluated. Biochar application significantly increased soil pH from 6.0 to nearly 7.0 and the available Mo content by 16–151
Radiocesium (137Cs) transfer from soil to crops is largely regulated by soil potassium (K) levels owing to the chemical similarity between K and cesium (Cs). However, the mitigation of Cs translocation in soybean through soil K is lower than in other crops, highlighting the importance of clarifying soybean-specific Cs translocation mechanisms. Although root nodule symbiosis has been proposed to alter nutrient transport systems, its impact on Cs dynamics remains unclear. We hypothesized that Cs translocation mechanisms are altered under root nodule symbiosis. To elucidate these mechanisms, we conducted field experiments using three soybean genotypes with different nodulation abilities and analyzed their elemental distribution patterns. Additionally, hydroponic experiments using inoculated soybeans were conducted to investigate 137Cs distribution. We found that Cs concentrations were consistently higher in nodules than in other organs. Radioisotope imaging also showed predominant 137Cs accumulation in nodules. Covariance analysis revealed that Cs translocation to shoot was lower in genotypes with higher nodule formation under the same soil exchangeable K conditions. Furthermore, increased nodule formation, especially nodule number, was associated with reduced Cs translocation to shoot. These results suggest that nodules contribute to suppressing Cs translocation to shoot and provide new insights into Cs dynamics under root nodule symbiosis.
It is known that sugar beet, derived from a halophytic wild ancestor, is highly salt-tolerant and that moderate concentrations of NaCl enhance its growth. However, the mechanisms of this improved growth performance by NaCl are not fully understood. In this study, we investigated the effects of NaCl on sugar beet growth under different light conditions and examined the relationship between light-induced oxidative stress and the beneficial effects of NaCl. Sugar beet seedlings in pots filled with fertilized soil were grown with different light intensities (low: 150-250 mu mol m-2s-1 or high: 600-1,000 mu mol m-2s-1) and NaCl treatments (irrigation with water with or without 50 mM NaCl) for 10 days. The enhanced sugar beet growth by NaCl application was only observed in plants with high light intensity treatments. Moreover, high light intensity treatment increased the shoot concentration of malondialdehyde, an indicator of oxidative stress, while NaCl application decreased it. While there was a positive correlation between Na concentration and the compatible solute glycine betaine in the shoots and roots under both light intensities, a significant negative correlation between the concentrations of glycine betaine and malondialdehyde was found only in the shoots under high light intensity. Because it is known that glycine betaine activates antioxidant enzyme activities and reduces oxidative stress responses, glycine betaine synthesized in response to NaCl accumulation may have reduced the oxidative stress caused by high light intensity in the shoots, which may have caused the enhanced growth in sugar beet. Furthermore, the growth promotion by NaCl accumulation was accompanied by a significant decrease in NO3-N concentration, which suggested that NaCl application affects nitrogen metabolism in sugar beet.
Since most leaf defense traits are derived from photosynthates, the increase of feeding damage in woody plants can be partially attributed to the recent increase in ground-surface O3 (elevated O3; eO3), which has suppressed the photosynthetic capacity of the plants. In different soil fertility conditions (fertile brown forest soil or infertile volcanic ash soil), we made research on leaf defense traits of the three species; birch (an early successional species), beech (a late successional species), and oak (an intermediate species). The results showed that except for lignin, the defensive capacity of birch leaves was reduced by eO3 as well as by a fertile soil condition. In beech as an O3-sensitive species, C/N was slightly higher in fertile soil; especially total phenolics and lignin concentrations, but increased with eO3. Therefore, carbon-nutrient balance hypothesis was not applicable in beech, which may be due to a different nutrient translocation system from birch and other trees. Oak, an O3-tolerant species, was less affected by eO3 and soil condition; C/N slightly tended to increase with eO3 due to a decrease in N. In addition, total phenolics of oak leaves tended to decrease by eO3 in infertile soil. From these results, O3 tolerance, which differs among tree species, can lead to new and altered plant-insect relationships through impact on defense functions, which can also lead to new relationships in biodiversity.
Cadmium (Cd) and arsenic (As) soil contamination poses significant problems for crop production worldwide. Zero-valent iron (Fe0) is a reactive material with reducing power capable of stabilizing toxic elements, including heavy metals and metalloids. In this study, we examined the effect of Fe0 application on Cd and As accumulation in spinach (Spinacia oleracea L.) grown in Cd- or As-contaminated soil. Fe0 soil application reduced Cd and As concentrations in spinach shoots and/or roots, and decreased their availability in soil, which may have affected their accumulation in spinach. Additionally, we examined the forms of soil Cd and As using the sequential extraction method. The potential availability of Cd and As forms in soils for plant root is: exchangeable > iron-manganese-oxide-bound > organic-matter-bound > free-oxide-occluded > residual. Fe0 application increased Cd and As distribution in the free-oxide-occluded fraction. Additionally, Fe0 application decreased As distribution in the iron-manganese-oxide-bound fraction while tended to decrease Cd in the organic-matter-bound fraction and increase it in the iron-manganese-oxide-bound fraction. Thus, Fe0 application can be effective in reducing Cd and As availability in their contaminated soils. Meanwhile, this study showed that Fe0 application inhibited the growth of spinach in Cd-contaminated soil. The unfavorable alterations caused by Fe0 in the contaminated soil should also be investigated in the future.
We assessed the effects of soil type on the concentrations of seven elements (calcium, potassium, sulfur, phosphorus, iron, manganese, zinc) in plants using x-ray fluorescence (XRF) analysis and plant specimens collected from calcareous and ultramafic soil areas, and 'general soils' (other types of soil represented by Brown Forest soils and Andosols) of Japan. Compared with the plants from other types of soils, the means of these seven elements were lower in plants originating from ultramafic soils, especially in phosphorus. Plants from calcareous soil areas had high iron and zinc concentrations on average, but this tendency was biased by plants collected on Mt. Fujiwara, which had extremely high average values of these elements. Calcium concentration in plants had a negative correlation with zinc or iron from calcareous or general soils, respectively, and iron showed positive correlations with zinc, manganese, and sulfur in plants from calcareous, ultramafic, and general soils, respectively. These correlations were not found in plants from general soils, suggesting that these relationships reflect the chemical characteristics of soil types. On the contrary, plants from all soil types showed a positive correlation between sulfur and phosphorus, whereas a positive correlation between potassium and phosphorus was found in plants from calcareous and general soils, but not in ultramafic plants, which instead showed a weak, but statistically significant correlation between potassium and sulfur. Statistically, iron and sulfur concentrations were positively correlated in all soil types, but manganese showed a low correlation with the other elements.
Palhinhaea cernua, a lycophyte, and Dicranopteris linearis, a fern, are commonly observed in solfatara fields in Kyushu, Japan, but their distribution trends are different. The aim of this study was to determine why P. cernua is more abundant in areas closer to fumaroles from both a soil and plant perspective. Samples of P. cernua and D. linearis, as well as their respective growing soils, were collected, and the mineral properties, including the concentration of various mineral elements and inorganic anions and δ15N, were determined. P. cernua was better adapted to soil with lower pH, higher soluble aluminum concentrations, and poorer calcium and phosphorus concentrations than D. linearis. A positive correlation was observed between shoot nitrogen concentration and both shoot sulfur concentration and soil water-soluble sulfur concentration in P. cernua, implying the involvement of sulfur in nitrogen acquisition in P. cernua. The results also suggested that D. linearis mainly uses soil NO3-N, while P. cernua uses NH4-N, which is predominant and excessive in the solfatara fields, particularly near the fumaroles. This high preference for NH4-N in P. cernua was confirmed through a cultivation experiment. While D. linearis prefers NO3-N and distributes further from fumaroles, P. cernua may have survived in the solfatara fields by utilizing NH4-N and sulfur, which are abundant near fumaroles where competition from other plant species is minimal.
Gypsum (calcium sulfate) are commonly used to improve subsoil acidity. Aluminum (Al) toxicity in acid soil reduces crop yields worldwide, especially in the tropical regions. In soil, sulfate decrease Al toxicity and improves plant growth and yield. Here, we aimed to investigate the effects of CaSO4 on Al stress of Al-tolerant accession Col-0 as well as the Al-sensitive accessions, Wei-0 and Ts-5, of in Arabidopsis thaliana using hydroponics. Our research indicates that CaSO4 supply has a significant effect on root growth and malate release compared to CaCl2 under Al stress, especially in Al-sensitive accessions. In response to Al stress, CaSO4 supply enhanced the expression of malate transporter gene AtALMT1 and sulfate transporter gene SULTR3;5, which were regulated by the Al-resistant transcription factor STOP1. Furthermore, knockout lines of SULTR3;5 and SULTR2;1, which are involved in sulfate uptake and translocation, were more Al-sensitive than the wild type under Al stress without CaSO4, and malate release was reduced with decreased AtALMT1 expression. In conclusion, one of the alleviative effects of gypsum application on Al rhizotoxicity is promoted by sulfate through organic acid release mechanism enhancing AtALMT1 expression, alongside the Ca2+ reduction Al3+ activity on the plasma membrane.
Andosols are soils characterized by relatively high sulfur concentrations, but also by their low phosphorus availability for crops and vegetables. We assessed the effect of soil type on the concentrations of sulfur and phosphorus in plants using handheld X-ray fluorescence (XRF) analysis on herbarium specimens collected from Andosols and non-volcanic soils in Japan. The sulfur concentrations in plants from Andosols were higher than plants from non-volcanic soils, but there was no significant difference between their foliar phosphorus concentrations. Only three specimens out of 1203 herbarium specimens were below the sulfur concentration (1000 mg kg-1) needed for adequate growth of crops, but about half of specimens were below that for phosphorus (2000 mg kg-1). The sulfur concentrations were high in herbaceous plants, especially in plants from the Brassicaceae and Liliaceae families. Overall, the results show that the sulfur concentrations in wild plants from Japan were higher in Andosols, with a low risk for sulfur deficiency. In contrast, the effect of soil type on the phosphorus concentrations in plants was small, and this may be explained because most wild plants grow under phosphorus-limiting conditions. Finally, the analysis reveals a strong correlation between sulfur and phosphorus concentrations in plants, suggesting the existence of a mechanism that controls the sulfur and phosphorus concentration ratios.