The role of phosphorus in growing tomatoes in near water-saturated soil

Abstract

High-yielding, waterlogged cultivations with considerable nitrogen input are widely practiced in Kyushu Island, Japan. Our objective was to determine the role of available phosphorus in relation to nitrogen and soil microorganisms on tomato growth in such systems. Tomato seedlings were grown in a pasteurized soil with ample KNO₃ in addition to different amounts of sodium or potassium phosphates to assess effects of phosphorus on growth enhancement and water-use efficiency in the seedlings grown with copious soil water (>－0.7?kPa). Both monobasic and dibasic phosphates applied at 40?mM to near-saturated soil markedly promoted seedling growth when fertilized with 120?mM KNO₃ or NH₄Cl, but not in the soil without nitrogen. The final concentration of NO₃^－ near the soil surface was maximized in the nitrated soil with no added PO₄^3－ but declined significantly when fertilized with any phosphates except Na₂HPO₄. No significant accumulation of NO₃^－ was detected in the water-saturated bottom soils regardless of soil fertilization with any plant minerals. Increased seedling growth was accompanied by lower dry root/shoot ratios and marked increases in evapotranspiration efficiency. Such positive effects of PO₄^3－ were greater with NO₃^－ than NH₄^＋ but diminished with increased concentrations of Na^＋ up to 160?mM. Nonetheless, growth promotions by ample NO₃^－ with PO₄^3－ were nearly negated in the non-pasteurized soil. Apparently, the activities of indigenous soil microbes were more significant than soil nitrogen and phosphorus in limiting the growth potential of tomato plants in near-saturated soils.     

Effects of soil redox potential (Eh) and pH on growth of sunflower and wheat

ABSTRACT

Soil redox potential (Eh) and pH are fundamentals parameters for plants growth. Measuring soil Eh is essential but complex due to the lack of measurement reliability resulting from high temporal variability and metrological challenges. This paper proposes practical advancements for measuring Eh in aerobic soils using combined electrodes (improvements in methodology specific to cleaning electrodes, measurement time, cleaning the dataset). The study of soil Eh and pH on sunflower and wheat in pot experiments has highlighted the relationship between Eh_cumul (Eh associated with a dimension of time) and the portion of porosity that is accessible to air. For reduced soil conditions, sunflower positively reacts to better aeration. Strong correlations exist between the duration of each potential range and the growth of sunflower. The study of sunflower growth in soil reveals extremely harmful impact resulting from too high and/or too low Eh values.     

Driving factors of soluble organic nitrogen dynamics in paddy soils: Structure equation modeling analysis

Soil soluble organic nitrogen (SON) is one of the most active components in soil nitrogen pools; however, limited information is available with regard to its driving factors, as well as their pathways and degrees of influence. In this study, structural equation modeling was used to analyze the driving factors, their significance, and pathways that affected SON dynamics in a waterlogged experiment of two typical paddy soils incubated for 80 d after green manure application. Soil pH, Eh, microbial biomass, enzyme activity, and SON dynamics were used to construct the structural equation model. Results showed that soil microbial biomass carbon (MBC), protease, glutamine, and initial organic matter (OM) directly and significantly affected soil SON with path coefficients corresponding to 0.405, 0.547, 0.523, and -0.623 (P < 0.01), respectively. Soil microbial biomass carbon and initial OM affected the SON dynamics indirectly through protease and glutamine activity. In addition, pH indirectly affected SON dynamics by glutamine activity. It is implied that soil MBC, protease, glutamine, and initial OM are the key factors affecting SON dynamics in the waterlogged paddy soils after green manure application. Our research indicated that structural equation modeling could provide an effective method to clearly recognize the impact, significance, and pathways of multiple factors on SON dynamics in paddy soils.     

The effects of planting systems on soil biology and quality attributes of tomatoes

This study was carried out to determine the effects of seedling types (grafted and non-grafted) and different plantation systems (raised-bed and flat planting) on growth, yield and quality of tomato (Solanum lycopersicum cv. Depar F1) plants organically grown in open-field conditions in Samsun. Soil microbial biomass-C increased by 25% and soil CO₂ contents increased by 16% in raised-bed systems. It was observed that soil microbial biomass-C positively correlated with CO₂ contents, leaf chlorophyll content, stomatal conductance and yield (P < 0.05 and P < 0.01) and negatively correlated with soil compaction. The highest chlorophyll content (47.37 CCI), fruit shape index (1.21) and yield (1.95 kg plant^?1) were obtained from grafted seedlings of the raised-bed planting systems. The highest stomatal conductance (118.50 mmol m^?2 s^?1) and firmness (79.34%) were obtained from non-grafted seedlings of the raised-bed planting systems. As a result, successful tomato cultivation was carried out with the use of raised-bed and grafted seedlings. However, non-grafted seedlings of the raised-beds had higher yield and quality values than the grafted seedlings of the flat planting.     

Methane and water soluble iron production under controlled soil pH and redox conditions

Abstract

When a soil is flooded, iron (Fe) reduction and methane (CH₄) production occurred in sequence as predicted by thermodynamics. The dissolution and precipitation of Fe reflected both soil pH and soil redox potential (Eh). The objective of our experiment was to determine both CH₄ production and Fe reduction as measured by Fe in solution in a flooded paddy soil over a wide range of closely controlled pH and Eh conditions. The greatest release of CH₄ gas occurred at neutral soil pH in combination with low soil redox potential (‐250 mV). Production of CH₄ decreased when soil pH was lowered in combination with an increase in the soil redox potential above ‐250 mV. Highest concentration of ferrous‐iron (Fe²⁺) under reducing conditions occurred when soil pH was lowered. Thus Fe reduction influenced CH4 formation in the flooded paddy soil. Results indicated that CH₄ production was inhibited by the process of ferric‐iron (Fe³⁺) reduction.     

Iodine desorption from rice paddy soil

Laboratory experiments on the desorption phenomena of iodine from rice paddy soil under waterlogged conditions, with a special reference to soil redox potential (Eh) and pH, have been conducted. Radioiodine tracer (1251), added to the soil, was readily sorbed on it. At the beginning of the waterlogging, the iodine desorption was low. However, iodine was desorbed into soil solution with time. The iodine desorption was enhanced markedly by the addition of organic substances such as straw pieces and glucose to the soil. Cultivation of rice plants in soil also affected the iodine desorption, suggesting root exudates and/or root autolysis might be participating in the desorption process. Eh dropped considerably after soil was waterlogged due to microbial metabolisms. Particularly low Eh values were observed in soils with plants and also with added organic substances. A negative correlation was seen between the desorption and soil Eh. High desorption was frequently observed when the Eh dropped to about -100 mV or below. Due to the reducing conditions (low Eh) by waterlogging, iodine in soil was leached into the soil solution; consequently total iodine concentration in paddy soil was considerably lower than forest and upland field soils. These iodine desorption phenomena under anaerobic conditions should be considered in assessing transfer of the long-lived radioiodine (129I) in the environment, especially in rice fields and marshland.     

Stability behavior of soil colloidal suspensions in relation to sequential reduction of soils

Flocculation and dispersion of colloidal particles of nine inorganic paddy soils were studied mainly based on turbidity measurements of the suspensions of soils which were previously incubated at 28°C under in vitro waterlogged conditions. After 1-week of incubation, the turbidity of the soils except for 1) two soils containing larger amounts of sodium salts and 2) one soil containing larger amounts of Fe and Al oxides, significantly decreased, and colloidal particles flocculated with 1) a decrease in soil Eh and 2) an increase in electric conductivity (EC). During the 3- to 4-week period of waterlogging, the turbidity of the three soils significantly increased with the 1) decrease in EC and 2) increase in pH of the soils although the Eh remained low. Infrared (IR) absorption analysis showed that the suspended colloidal particles consisted of layer silicates from respective soil clays. Oxidation of suspensions of waterlogged soils by air-bubbling led to an increase in turbidity with the 1) increase in Eh, and 2) decrease in pH, EC, and water-soluble Fe₂₊ concentration. It was suggested that the stability of the soil colloidal suspensions was affected by soil reduction with alterations in ionic species and their concentrations at clay surfaces and in soil solutions.     

Saccharidic compounds as soil redox effectors and their influence on potential N2O production

Cellulose, xylan, and glucose were compared in waterlogged soil as modifying factors of the redox potential (Eh), of the quantity of reducing equivalents, and of the soil capacity to produce N₂O and CO₂. During the study period (168 h) soils supplied with glucose and xylan showed a higher Eh decrease than the control soil and the soil treated with cellulose. In samples taken after 0, 24, 48, and 168 h, the soils supplied with C showed a higher number of reducing equivalents than the control soil did. These quantities were not correlated with Eh values, nor with N₂O production. N₂O production was increased compared with the control soil over the entire experimental period in the glucose-amended soils but only after 48 h in the xylan-amended soils and not until 168 h in the cellulose-treated soils. The CO₂:N₂O ratio was consistently higher than the theoretical value of 2, suggesting that denitrification and CO₂ production via fermentation occurred simultaneously. Moreover, this ratio was highly correlated with the Eh values. We conclude that more research is needed to explain the role of soil redox intensity (Eh) and capacity (quantity of redox species undergoing reduction) in the expression of soil denitrification-fermentation pathways.     

Ozone,acidic precipitation,and soil Mg impacts on soil and loblolly pine seedling nutrient status after three growing seasons

Recent studies have suggested that the growth of loblolly pine (Pinus taeda L.) has declined in the southeastern United States, possibly due to acidic deposition and air pollutants, especially under conditions of low nutrient availability. Consequently, the potential for individual and synergistic impacts of O₃, acidic precipitation, and soil Mg status on the nutrient status of loblolly pine seedlings and soil was investigated over a 3 yr study period. Thirty-six open top chambers equipped with a rainfall exclusion/addition system were utilized to administer three levels of O₃ (subambient, ambient, or twice ambient) and two acidic precipitation treatments (pH 3.8 or 5.2) to seedlings growing in 24-L plastic pots containing soil having either 35 or 15 mg kg^?1 of exchangeable Mg. Each chamber contained 36 pots, and each treatment combination was replicated six times for a total of 1296 individual pots. After three seasons, throughfall and foliar nutrition data indicated that foliar leaching was not accelerated by increasing the acidity of precipitation from pH 5.2 to 3.8 and that increasing O₃ did not act to exacerbate foliar leaching. Further, foliar nutrient concentrations were not significantly affected by precipitation pH or O₃ treatments. Soil and soil solution data also indicate no accelerated soil leaching associated with chronic acidic precipitation. Differences in soil Mg treatments were reflected in soil solution and seedling Mg contents, but the 15 mg kg^?1 soil Mg treatment was not sufficiently low enough to induce Mg deficiency in the seedlings.     

Nitrogen Inputs with Different Substrate Quality Modified pH,Eh, and N Dynamics of a Paddy Soil Incubated under Waterlogged Conditions

Synthetic fertilizer, livestock manure, and green manure are the typical nitrogen (N) sources in agriculture. This study was conducted to investigate the effects of different N sources on soil chemical environment and N dynamics. Changes in pH, redox potential (Eh), and concentration and δ¹⁵N of dissolved N [ammonium (NH₄⁺), nitrate (NO₃^?), organic N, and total N] of soils treated with urea (U), pig manure compost (PMC), and hairy vetch (HV) were investigated in an incubation experiment under waterlogged conditions. The patterns of pH, Eh, and N concentration reflected both a greater mineralization potential of N derived from U than that from HV and PMC and easier decomposability of HV than PMC. The δ¹⁵N further suggested that nitrification was more active for U than for HV- and PMC-treated soils and that N loss via NH₃ volatilization and denitrification would be greater for HV than U and PMC treatments.     

Factors controlling carbon dioxide and methane production in acid sulfate soils

Methane and C02 production in flooded acid sulfate soils of Thailand were governed primarily by soil oxidation-reduction potential (Eh) and pH. The critical Eh and pH levels at which CH₄ emission began was Eh-150 mV, and pH 6.1. Low soil pH limited soil reduction and subsequently CH₄ production. Soil respiration (C02 production) was influenced by Eh-pH levels and organic matter content. Soils with higher C02 production rates produced greater amounts of CH₄. Soil pH, however, was the dominant variable which influenced organic matter decomposition, low soil Eh conditions and subsequent CH₄ and CO₂ production. Curvilinear or log transformations of pH, Eh and organic matter content (OM) were used in explaining variables controlling CH₄ and CO₂ production; CH₄ = ?2.359 ? 0.0001 Eh + 2.047 pH ? 3.019 (In pH)² CO₂ = ?5210 ? 1.6 Eh + 3144 (In pH) + 1011 (In OM).     

Influence of nitrate on methane production and oxidation in flooded soil

Abstract

Methane (CH₄) production in paddy soils and sediments as influenced by nitrate (NO₃) addition was studied under a closely controlled soil pH and oxidation‐reduction (redox) potential (Eh) conditions. CH₄ production was affected by soil pH and NO₃ ^‐. Added NO₃ ^‐ reduced the amount of CH₄ produced of each pH level studied. Nitrate addition primary effect in reducing CH₄ production was through the resultant increase in soil redox potential (Eh). Using Methyl Fluoride (MF), a CH₄ oxidation inhibitor we found that added NO₃ ^‐ was not used in CH₄ oxidation by methanotrophic bacteria.     

An investigation of the impact of Glomus clarum (mycorrhiza) on the growth of tomato (Lycopersicum esculentum mill.) on both sterilized and non-sterilized soils

Abstract

A greenhouse experiment was carried out to investigate the influence of Glomus clarum (mycorrhiza) on the growth of tomato seedlings grown in both sterilized and non-sterilized soils. Highest growth parameter values were recorded in tomato plants inoculated with mycorrhiza but grown in sterilized soil, followed by those grown in non-sterilized soil but inoculated with mycorrhiza also. Sterilized but non-inoculated tomato plants also had growth and were closely followed by non-sterilized, non-inoculated tomato plants. There was no significant difference in all the treatments when girth of the tomato plants used was measured in this study. Nutrient uptake (N,P,K) was significantly found highest in the inoculated sterilized tomato plants while it was found lowest in the non-sterilized, non-inoculated tomato plants. Generally, mycorrhizal-inoculated tomato plants (whether sterilized or non-sterilized) showed better growth in all the treatments used.     

Influence of pH on the redox chemistry of metal (hydr)oxides and organic matter in paddy soils

Purpose

The primary purpose of this study was to determine how flooding and draining cycles affect the redox chemistry of metal (hydr)oxides and organic matter in paddy soils and how the pH influences these processes. Our secondary purpose was to determine to what extent a geochemical thermodynamic equilibrium model can be used to predict the solubility of Mn and Fe during flooding and draining cycles in paddy soils.

Material and methods

We performed a carefully designed column experiment with two paddy soils with similar soil properties but contrasting pH. We monitored the redox potential (Eh) continuously and took soil solution samples regularly at four depths along the soil profile during two successive flooding and drainage cycles. To determine dominant mineral phases of Mn and Fe under equilibrium conditions, stability diagrams of Mn and Fe were constructed as a function of Eh and pH. Geochemical equilibrium model calculations were performed to identify Mn and Fe solubility-controlling minerals and to compare predicted total dissolved concentrations with their measured values.

Results and discussion

Flooding led to strong Eh gradients in the columns of both soils. In the acidic soil, pH increased with decreasing Eh and vice versa, whereas pH in the alkaline soil was buffered by CaCO₃. In the acidic soil, Mn and Fe solubility increased during flooding due to reductive dissolution of their (hydr)oxides and decreased during drainage because of re-oxidation. In the alkaline soil, Mn and Fe solubility did not increase during flooding due to Mn(II) and Fe(II) precipitation as MnCO₃, FeCO₃, and FeS. The predicted levels of soluble Mn and Fe in the acidic soil were much higher than their measured values, but predictions and measurements were rather similar in the alkaline soil. This difference is likely due to kinetically limited reductive dissolution of Mn and Fe (hydr)oxides in the acidic soil. During flooding, the solubility of dissolved organic matter increased in both soils, probably because of reductive dissolution of Fe (hydr)oxides and the observed increase in pH.

Conclusions

Under alternating flooding and draining conditions, the pH greatly affected Mn and Fe solubility via influencing either reductive dissolution or carbonate formation. Comparison between measurements and geochemical equilibrium model predictions revealed that reductive dissolution of Mn and Fe (hydr)oxides was kinetically limited in the acidic soil. Therefore, when applying such models to systems with changing redox conditions, such rate-limiting reactions should be parameterized and implemented to enable more accurate predictions of Mn and Fe solubility.     

旱改水对水稻幼苗生长的影响及秸秆的改良作用

本研究以江汉平原旱改水为研究背景,采用土壤盆栽试验和室内淹水培养相结合的方法,以多年水稻土为对照,研究了多年棉田土旱改水及添加秸秆(9 g·kg-1)对水稻幼苗生长和矿质元素吸收的影响以及土壤氧化还原电位和有效态铁、锰、铜、锌含量变化,为旱改水水稻的种植提供参考。结果表明,棉田土旱改水后,水稻幼苗生长缓慢并出现失绿黄化症状,其地上部干重和叶绿素含量仅分别约为水稻土处理的30%和20%。旱改水处理水稻植株Fe含量显著低于、而Cu和Zn含量则显著高于水稻土处理。棉田土旱改水土壤氧化还原电位(Eh)显著高于水稻土;淹水处理10 d,土壤DTPA-Fe含量仅为水稻土的7%左右,而DTPA-Cu和DTPA-Zn含量则分别是水稻土的1.4~2.5倍和1.6~1.8倍。随着淹水时间的延长,棉田土旱改水土壤有效态铁含量逐渐增加,有效态锰、铜和锌含量呈先升高后降低趋势;到淹水处理的第28 d,棉田土旱改水土壤有效态铁、锰、铜和锌含量与水稻土之间的差异逐渐缩小。Fe不足及Cu过量可能是导致旱改水水稻幼苗生长缓慢、失绿黄化的主要原因。旱改水条件下添加秸秆可以降低土壤的Eh值,提高土壤DTPA-Fe含量及降低土壤DTPA-Cu和DTPA-Zn含量,显著提高旱改水初期水稻幼苗叶绿素含量,但对水稻生物量无显著影响。添加秸秆并不能完全消除旱改水对水稻幼苗生长的抑制作用。     

Silicon concentrations in soil and bark in Irish Sitka spruce forests

The relationship between plant‐available silicon (Si) soil concentrations and bark Si concentrations in coniferous species is poorly understood. The objectives of this research were to generate baseline data on Si concentrations in soils and bark of Sitka spruce (Picea sitchensis) seedlings in Ireland and to understand better the relationship between soil and bark Si concentrations. Seedlings were harvested from eight plantation forestry sites and two tree nurseries, and Si concentrations in the bark tissue as well as plant‐available Si concentrations in soils (CaCl₂ extractant) were measured. Bark Si concentrations varied significantly between sites and were lowest [mean 790 (± 242 SD) mg kg^?1 dry plant tissue] on acidic, organic rich peat soils, while the highest Si concentrations occurred in seedlings [mean 3688 (± 633 SD) mg kg^?1 dry plant tissue] grown on soils with low C concentration and higher pH values (≈ 4.5 to 5.5 in H₂O). Plant‐available Si soil concentrations were not related to soil C concentrations. There was a negative (but statistically not significant) relationship between plant–soil concentrations and soil pH. A significant negative relationship was observed between plant‐available soil Si concentrations and bark Si concentrations, which may be related to the presence of soil from mixed soil horizons forming the mounds that seedlings were planted on. Uptake and sequestration of Si by seedlings may have been related to the rate of growth of the seedlings, as bark Si concentrations were highest on sites that were expected to have greater seedling growth rates. The negative relationship between bark and plant‐available Si soil concentrations suggest that uptake of Si by Sitka spruce is rejective at higher concentrations.     

Relationship between soil chemical properties and microbial metabolic patterns in intensive greenhouse tomato production systems

ABSTRACT

Intensive greenhouse production involving excessive fertilizer and organic manure application rates may affect soil chemical and biological quality. Soil samples from 50 commercial greenhouses for tomato production in northern China were collected for the evaluation of the status of soil fertility and identification of the soil chemical factor that exerts the strongest influence on microbial functional diversity. The soil total nitrogen content showed high soil fertility and was 68% higher than 1000 mg kg^?1 and 14% higher than 1500 mg kg^?1. Differential soil pH values caused statistically significant shifts in microbial metabolic activity (average well color development, AWCD) and Shannon’s diversity index using Biolog^TM ECO plates assay. The highest soil microbial functional diversity was observed at near neutral pH values. When individual data points were plotted against soil organic matter (SOM), significant positive associations with soil microbial biomass nitrogen and AWCD were observed. The canonical correspondence analysis confirmed that shifts in the soil microbial functional diversity were associated with changes in pH, total nitrogen, and SOM. This study indicated that excessive fertilization changed the community-level physiological profile of the soil microorganisms, and this effect can be a consequence of changes in soil pH under intensive greenhouse management.     

Properties and metabolic diversity of microbial communities in soils treated with steam sterilization compared with methyl bromide and chloropicrin fumigations

The effects of steam sterilization (SS) on soil microbial properties including metabolic diversity of the microbial communities were examined in a greenhouse compared with those of two fumigants, methyl bromide (MB) and chloropicrin (CP). The numbers of fungi decreased in all the treatments. Nitrifiers, both ammonium-oxidizing bacteria and nitrite-oxidizing bacteria, were severely affected by the SS and CP treatments, resulting in their virtual disappearance. The decrease in the levels of microbial biomass C and N after the treatments suggested that the SS and CP treatments eradicated the microorganisms more effectively than the MB treatment and that the influence of the former persisted until the end of the experiment, 4 months after the treatments. Accumulation of NH₄-N was observed after the SS and CP treatments mainly due to the partial decomposition of the dead microorganisms and the marked decrease in the number of ammonium-oxidizing bacteria. The richness and average well color development (AWCD) values in the microbial communities after the SS treatment were evaluated by the carbon substrate utilization method using Biolog ECO MicroPlates. The values decreased markedly immediately after the treatment but showed a rapid recovery, while those after the CP treatment continued to decrease until the transplanting of tomato seedlings. The effect of the MB treatment on the soil microbial communities was less pronounced. The growth of the tomato plants was promoted by the SS and CP treatments due to the increase in the N supply at the initial stage of tomato growth.     

Effect of phosphate rock, lime and cellulose on soil microbial biomass in acidic forest soil and its significance in carbon cycling

Phosphate rock (PR), limestone, coal combustion by-product (CCBP) high in Ca and high organic manures are potential amendments for increasing agricultural production in the acidic soils of the Appalachian region. The objective of this study was to examine effects of PR, CCBP and cellulose addition on soil microbial biomass in an acidic soil based on the measurement of soil microbial biomass P (P _mic) and on the mineralization of organic matter. Application of PR alone or in combination with CCBP increased P _mic. The P _mic was far less when the soil received PR in combination with limestone than with PR application alone or PR in combination with CCBP. Either CCBP or limestone application alone considerably decreased P _mic in the soil due to reduced P solubility. Cellulose addition alone did not increase P _mic, but P _mic was significantly increased when the soil was amended with cellulose in combination with PR. The decomposition of added cellulose was very slow in the soil without PR amendment. However, mineralization of both native organic matter and added cellulose was enhanced by PR application. Mineralization of organic matter was less when the soil was amended with PR in combination with high rates of CCBP (> 2.5%) because PR dissolution varied inversely with amount of CCBP addition. Overall, CCBP had no detrimental effect on soil microbial biomass at low application rates, although, like limestone, CCBP at a high rate may decrease P _mic in P-deficient soils through its influence on increased soil pH and decreased P bioavailability in the soil. Application of PR to an acidic soil considerably enhanced the microbial activity, thereby promoting the cycling of carbon and other nutrients. Received: 11 December 1995     

Seasonal dynamics of soil microbial biomass C shows close correlation with environmental factors in natural Fagus crenata forests

Abstract

Soil microbial biomass (C_mic) is an important factor regulating a number of ecosystem processes. In this study, we investigated seasonal variations in soil microbial biomass in natural climax beech (Fagus crenata) forests in a typical cold-temperate mountain region of Japan. Four permanent tower sites along an altitudinal gradient were selected and soil samples were collected once every month during the growing season of 2007. Soil microbial biomass (by fumigation-extraction method) and soil properties were later measured in the laboratory, while environmental factors (soil temperature, soil moisture) were continuously recorded in the field. Our results indicated large seasonal variations (130.4 ~ 5558.0 µg g^?1) in soil microbial biomass in beech forests – a range that is much larger than previously reported. Statistically significant correlations are noted between soil properties with C_mic, but largely due to spatial linkages. On the other hand, the environmental factors of soil temperature and especially soil moisture largely control seasonal variations in C_mic. Furthermore, pH could be an important factor influencing seasonal change in C_mic at the 20–30 cm deep soil layer. The study suggests no direct correlation between plant eco-physiology and soil microbial biomass in seasonal courses of the forests.