Influence of soil properties on coastal sandplain grassland establishment on former agricultural fields

The decline in species‐rich grasslands across the United States has increased the importance of conservation and restoration efforts to preserve the biodiversity supported by these habitats. Abandoned agricultural fields often provide practical locations for the reestablishment of species‐rich grasslands. However, these fields often retain legacies of agriculture both in their soils, which may have higher pH and nitrogen (N) contents than soils that were never farmed, and in their plant communities, which are dominated by non‐native species and poor in native seed stock. We considered methods of reversing these legacies to create native‐species‐rich grassland on former agricultural land. We tested seeding and tilling combined with additions of sulfur (S), carbon (C), N or water to establish diverse sandplain grassland vegetation on an old field on Martha's Vineyard, Massachusetts. We measured soil pH, extractable nitrate and ammonium, and total and native species richness and native species cover for 5 years after treatment. S additions lowered pH to values typical of never‐tilled sandplain ecosystems and increased native species cover, but had no effect on species richness. C, N, and water additions had no significant effects on the soil or vegetation. Seeding and tilling were more effective at restoring native species richness than any soil amendments and indicated a greater importance of biotic factors compared with soil conditions in promoting sandplain vegetation establishment. S amendment accelerated establishment of native species cover for several years but the effect of S additions compared with seeding and tilling alone declined over time.     

Effect of Sheep Urine Deposition on the Bacterial Community Structure in an Acidic Upland Grassland Soil

The effect of the addition of synthetic sheep urine (SSU) and plant species on the bacterial community composition of upland acidic grasslands was studied using a microcosm approach. Low, medium, and high concentrations of SSU were applied to pots containing plant species typical of both unimproved (Agrostis capillaris) and agriculturally improved (Lolium perenne) grasslands, and harvests were carried out 10 days and 50 days after the addition of SSU. SSU application significantly increased both soil pH (P < 0.005), with pH values ranging from pH 5.4 (zero SSU) to pH 6.4 (high SSU), and microbial activity (P < 0.005), with treatment with medium and high levels of SSU displaying significantly higher microbial activity (triphenylformazan dehydrogenase activity) than treatment of soil with zero or low concentrations of SSU. Microbial biomass, however, was not significantly altered by any of the SSU applications. Plant species alone had no effect on microbial biomass or activity. Bacterial community structure was profiled using bacterial automated ribosomal intergenic spacer analysis. Multidimensional scaling plots indicated that applications of high concentrations of SSU significantly altered the bacterial community composition in the presence of plant species but at different times: 10 days after application of high concentrations of SSU, the bacterial community composition of L. perenne-planted soils differed significantly from those of any other soils, whereas in the case of A. capillaris-planted soils, the bacterial community composition was different 50 days after treatment with high concentrations of SSU. Canonical correspondence analysis also highlighted the importance of interactions between SSU addition, plant species, and time in the bacterial community structure. This study has shown that the response of plants and bacterial communities to sheep urine deposition in grasslands is dependent on both the grass species present and the concentration of SSU applied, which may have important ecological consequences for agricultural grasslands.     

The role of ant nest-mounds in maintaining small-scale patchiness in dry grasslands in Central Germany

Nest-mounds of Lasius flavus, Lasius alienus and Formica rufibarbis are mainly on north-facing slopes with a markedly patchy distribution at Gimritz, northwest of Halle (Saale). Nest-mound soils had higher sodium and potassium levels but lower phosphorus and nitrogen levels than soils away from mounds. Moisture content was lower but soil pH was significantly higher in nest-mound soils. However, there was no difference in the relative abundance of plants of alkaline or acid soils growing on mounds compared to plants off mounds and no significant difference between the numbers of nitrophilous plants on and off mounds. Fewer plant species (35 spp.) in total occurred on mounds than off mounds (42 spp.). Plant species richness and total plant cover was significantly less on the nest-mounds than off the mounds. Similarly, the mean number of grass (1.39 spp dm−2) and forb species (1.13 spp dm−2) on mounds was lower than the mean number of grass (1.99 spp dm−2) and forb species (1.91 spp dm−2) off mounds. Some shrubs and forbs had higher occurrence and cover on the mounds, although only in Calluna vulgaris, Thymus serpyllum and Cerastium arvense was this difference particularly marked. Ant nest-mounds may favour plant species that cannot compete with tall grasses on unmanaged, set-aside meadows. This revised version was published online in November 2006 with corrections to the Cover Date.     

Relation of the snow cover to the structure of vegatation in the alpine communities of the eastern Tsinghai-Tibet Plateau

The type of snow cover considerably influences the sctructure of vegetation and production-related processes in alpine communities of diverse regions. The relation of snow cover thickness to the structure of apline plant communities in the eastern Tsinghai-Tibet Plateau (Sichuan, People's Republic of China) was studied by analyzing the vegetation in 251 sample areas grouped in five transects along the gradient of mesotopographic conditions and wintertime snow cover thickness. Considerable differentiation of plant communities related to snow cover thickness in revealed along the line from the northern to the southern slope. It is shown that the influence of snow cover on the distribution of particular plant species is significantly greater than the influence of soil properties (pH, content of humus, total phosphorus, potassium, and ammonia nitrogen in the upper strata of soil). Among the 56 herbaceous plant species and 7 shrub species studied, 52 herbaceous and all the 7 shrub species showed significant (P < 0.05) correlation (positive or negative) to snow cover thickness. Snow cover thickness appeared also significantly correlated to a number of soil properties: soil thickness, content of water, total phosphorus, and humus. But, in contrast to the alpine communities of the Caucasus and the Alps, no considerable acidification of the soil under snowflakes, due to perpetual removal of cations by melt water, was observed.     

The Effects of Frost-Heaving on Objects in Soils

Abstract

Frost-heaving of natural and cultural materials occurs in soils whenever soil temperatures drop below freezing and the soils are moist. Frost-heaving is greatest in imperfectly drained, silt-textured, porous soils, at shallow depths, when rate of frost penetration is slow and when no vegetation or snow cover is present. The density, size, shape, orientation, and thermal properties of materials also affect the rate of frost-heaving. Because the movement is accumulative, the longer an object is buried, the greater will be its displacement. Frost-heave is a process which can mix stratigraphy and cultural materials at archaeological sites.     

Size, activity and catabolic diversity of the soil microbial biomass in a wetland complex invaded by reed canary grass

Reed canary grass (Phalaris arundinacea, L.) invasion of wetlands is an ecological issue that has received attention, but its impact on soil microbial diversity is not well documented. The present study assessed the size (substrate-induced respiration), catabolic diversity (CLPP, community level physiological profiles) and composition (selective inhibition) of the soil microbial community in invaded (>95% P. arundinacea cover) and in non-invaded areas of a wetland occupied by native species grown either as a mixed assemblage (22 species) or as quasi-monotypic stands of Scirpus cyperinus (74% cover). The study also tested the hypothesis that decomposition of lignin- and phenolics-rich plant tissues would be fastest in soils exhibiting high catabolic diversity. Results showed that soil respiration, microbial biomass and diversity were significantly higher (P?<?0.03; 1.5 to 3 fold) in P. arundinacea-invaded soils than in soils supporting native plant species. Fungal to bacterial ratios were also higher in invaded (0.6) than in non-invaded (0.4) plots. Further, canonical discriminant analysis of CLPP data showed distinct communities of soil decomposers associated with each plant community. However, these differences in microbial attributes had no effect on decomposition of plant biomass which was primarily controlled by its chemical composition. While P. arundinacea invasion has substantially reduced plant diversity, this study found no parallel decline in the size and diversity of the soil microbial community in the invaded areas.     

Plant communities and plant production in the western Queen Elizabeth Islands

A study of soils, plant communities, and net annual plant production was conducted with 41 stands at 3 sites on 3 arctic islands. Twelve additional sites were studied in less detail on Ellef Ringnes, King Christian and Melville islands and on four other islands. Through polar ordination five groupings were recognized. Alopecurus and Puccinellia barrens on sand to silty soils and on silty soils, high in sodium salts respectively. Species richness averaged 2.6 ± 2.0 and total plant cover 6.8 ± 2.7%. The Phippsia barrens occur on sheet eroded surfaces and in gulleys with deep winter snow. Species richness was 9.8 ± 5.0 and total plant cover 14.8 ± 9.6%. The graminoid steppes on sandy soils averaged 7.6 ± 2.4 species and total plant cover 40.0 ± 2.8%. Eight stands were dominated by moss-graminoids, mostly on loam soils. Species richness was 24.9 ± 3.4 and total plant cover 77.7 ± 16.1%. Plant producion was 8.0 g m⁻² in a Puccinellia barren and 9.4 g m⁻² in a Luzula confusa graminoid steppe. Net annual production ranged from 18.8 to 58.7 in 6 other stands. The 13 stands within the cryptogam-herb community complex occur on sandy loam to clay-loam soils. Species richness averaged 26.3 ± 6.2 and total plant cover 61.2 ± 24.7%.
Mosses and lichens play a significant role in the establishment and maintenance of communities with a greater species richness and plant production of vascular plant species. The ability of mosses to hold moisture and the presence of limited bluegreen algae that fix nitrogen appear essential to the maintenance of greater species richness, plant cover and plant production compared with the barren polar deserts that are often nearby.     

Testate amoebae and environmental features of polygon tundra in the Indigirka lowland (East Siberia)

Polygon tundra characterizes large areas of arctic lowlands. The micro-relief pattern within polygons offers differentiated habitats for testate amoeba (testacean) communities. The objective of this study was to relate testacean species distribution within a polygon to the environmental setting. Therefore, testaceans from four cryosol pits dug at different locations within a low-centered polygon were studied in the context of pedological and pedochemical data, while ground temperature and ground moisture were measured over one summer season. The study site is located on the Berelekh River floodplain (Indigirka lowland, East Siberia). The environmental data sets reflect variations along the rim-to-center transect of the polygon and in different horizons of each pit. The testacean species distribution is mainly controlled by the soil moisture regime and pH. Most of the identified testaceans are cosmopolitans; eight species are described from an arctic environment for the first time. Differences in environmental conditions are controlled by the micro-relief of polygon tundra and must be considered in arctic lowland testacean research because they bias species composition and any further (paleo-)ecological interpretation.     

Effect of sheep urine deposition on the bacterial community structure in an acidic upland grassland soil

The effect of the addition of synthetic sheep urine (SSU) and plant species on the bacterial community composition of upland acidic grasslands was studied using a microcosm approach. Low, medium, and high concentrations of SSU were applied to pots containing plant species typical of both unimproved (Agrostis capillaris) and agriculturally improved (Lolium perenne) grasslands, and harvests were carried out 10 days and 50 days after the addition of SSU. SSU application significantly increased both soil pH (P < 0.005), with pH values ranging from pH 5.4 (zero SSU) to pH 6.4 (high SSU), and microbial activity (P < 0.005), with treatment with medium and high levels of SSU displaying significantly higher microbial activity (triphenylformazan dehydrogenase activity) than treatment of soil with zero or low concentrations of SSU. Microbial biomass, however, was not significantly altered by any of the SSU applications. Plant species alone had no effect on microbial biomass or activity. Bacterial community structure was profiled using bacterial automated ribosomal intergenic spacer analysis. Multidimensional scaling plots indicated that applications of high concentrations of SSU significantly altered the bacterial community composition in the presence of plant species but at different times: 10 days after application of high concentrations of SSU, the bacterial community composition of L. perenne-planted soils differed significantly from those of any other soils, whereas in the case of A. capillaris-planted soils, the bacterial community composition was different 50 days after treatment with high concentrations of SSU. Canonical correspondence analysis also highlighted the importance of interactions between SSU addition, plant species, and time in the bacterial community structure. This study has shown that the response of plants and bacterial communities to sheep urine deposition in grasslands is dependent on both the grass species present and the concentration of SSU applied, which may have important ecological consequences for agricultural grasslands.     

Competitive dominance mediates the effects of topography on plant richness in a mountain grassland

Small-scale landforms influence plant species richness, but their mechanisms and effects in semi-natural dry grasslands have been poorly investigated. In this study we compared vascular plant richness, species composition, plant traits, soil properties and biomass nutrient content of convex (hillocks) and concave (hollows) karst landforms in a mountain pasture of the Central Apennines (Italy), at a small spatial scale (1 m² plots). We found hillocks had significantly higher species richness than hollows. On hillocks, smaller Specific Leaf Area and Lateral Width, together with greater allocation of resources to Below-Ground Organs, indicated lower water availability, whereas hollows had deeper (thus moister), more acidic and more fertile soils, with aboveground plant biomass displaying higher nutrient levels. Partial correlation and regression tree models suggested that fine-scale richness patterns were not directly determined by abiotic properties, but were rather the result of competition levels associated with the cover of Agrostis capillaris (=A. tenuis) – a calcifuge and drought-sensitive grass able to achieve dominance only in hollows. The higher functional convergence exhibited by hollows suggests that A. capillaris is a strong competitor both above- and below-ground, mediating the effects of topography by imposing a biotic filter. On hillocks, competition is released by lower levels of available soil water in summer and higher soil pH, resulting in higher species richness and a more functionally divergent assemblage.     

Phosphorus pools and other soil properties in the rhizosphere of wheat and legumes growing in three soils in monoculture or as a mixture of wheat and legume

Background and aims

Phosphorus and nitrogen availability and forms are affected by soil properties as well as by plant species and further modulated by soil microbes. Additionally, close contact of the roots of two plant species may affect concentrations and forms of N and P. The aim of this study was to assess properties related to N and P cycling in the rhizosphere of wheat and legumes grown in monoculture or in wheat/legume mixtures in three soils differing in pH.

Methods

Faba bean, white lupin and wheat were grown in three soils differing in pH (4.8, 7.5 and 8.8) in monoculture or in mixed culture of wheat and legumes. Rhizosphere soil was collected at flowering and analyzed for P pools by sequential fractionation, available N as well as community structure of bacteria, fungi, ammonia oxidizers, N₂-fixers and P mobilizers by polymerase chain reaction (PCR)—denaturing gradient gel electrophoresis (DGGE).

Results

Soil type was the major factor determining plant growth, rhizosphere nutrient dynamics and microbial community structure. Among the crop species, only faba bean had a significant effect on nitrification potential activity (PNA) in all three soils with lower activity compared to the unplanted soil. Soil type and plant spieces affected the community composition of ammonia-oxidizing archaea (AOB), ammonia-oxidizing archaea (AOA), N₂-fixers (nifH), P mobilizers (ALP gene) and fungi, but not that of bacteria. Among the microbial groups, the AOA and nifH community composition were most strongly affected by crop species, cropping system and soil type, suggesting that these groups are quite sensitive to environmental conditions. All plants depleted some labile as well as non-labile P pools whereas the less labile organic P pools (NaOH extractable P pools, acid extractable P pools) accumulated in the rhizosphere of legumes. The pattern of depletion and accumulation of some P pools differed between monoculture and mixed culture as well as among soils.

Conclusions

Plant growth and rhizosphere properties were mainly affected by soil type, but also by crop species whereas cropping system had the least effect. Wheat and the legumes depleted less labile inorganic P pools in some soils whereas less labile organic P pools (NaOH extractable P, acid extractable P) accumulated in the rhizosphere of legumes.     

A plant diversity×water chemistry experiment in subalpine grassland

Plant diversity has been shown to drive important ecosystem functions such as productivity. At the same time, plant diversity and species composition are altered in alpine ecosystems by human impacts such as skiing. Therefore, we investigated impacts of decreased species richness and ski piste treatments on ecosystem functions in subalpine grassland.Species richness manipulations were combined with nutrient input from snow cover treated with snow additives that are commonly used on ski pistes. Three different species richness levels containing 1, 3 or 9 species randomly selected from a larger pool plus unmanipulated meadow plots were treated with four water types to simulate melt water. One water type contained the snow additive ammonium nitrate. Invasion into the communities was prevented by weeding during 2 years and allowed in three subsequent years.Higher species richness increased plant cover and biomass and decreased their variation. The number of functional groups in a plant assemblage had a positive effect on plant growth. Ammonium nitrate strongly increased biomass and plant cover after a single application but decreased species richness in originally diverse meadow plots. There was no significant interaction between species richness and water-type treatments.After the cessation of weeding, the species richness of different plot types converged within 3 years due to invasion. Nevertheless, relationships between initial species richness and plant cover remained positive.The results suggest that the diversity and species composition of alpine vegetation are important factors influencing cover and biomass, in particular during re-colonization of bare ground after disturbances such as ski-piste construction. In slow-growing alpine vegetation, initially positive diversity effects may remain even after successional convergence of species richness due to invasion. The negative effect of ammonium nitrate on species richness suggests the snow additives should only be used with care.     

Factors influencing macroinvertebrate colonization of seasonal wetlands: responses to emergent plant cover

1. We conducted field experiments to examine factors influencing macroinvertebrate colonization of seasonally flooded marshes. Few macroinvertebrate species were found aestivating in soils within non-flooded wetlands indicating that most taxa colonize these marshes from other flooded habitats.
2. We manipulated amounts of salt grass ( Distichlis spicata ) to examine how emergent plant cover affects aerial colonization by macroinvertebrates. Areas mowed 3 weeks before flooding had low plant cover, areas mowed 5 and 9 weeks before flooding had medium and high plant cover, respectively, and non-mowed control areas had the most plant cover. Macroinvertebrate numbers and biomass were generally higher in mowed treatment areas than in control areas, but overall diversity was generally higher in high plant cover and control areas than in low plant cover areas.
3. Mosquitoes (Culicidae), brine flies (Ephydridae) and hover flies (Syrphidae) were positively correlated with amount of plant cover, and waterboatmen (Corixidae), midges (Chironomidae) and water scavenger beetles (Hydrophilidae) were negatively correlated with plant cover. Species assemblages changed seasonally among treatment areas because these taxa colonize wetlands at different times in the year.
4. These results demonstrate that invertebrate communities may be different within plant stands with heterogeneous amounts of emergent cover, and management practices that alter the structure of wetland vegetation can influence macroinvertebrate communities colonizing seasonal marshes.     

Differences in Earthworm Densities and Nitrogen Dynamics in Soils Under Exotic and Native Plant Species

Previous studies of the invasion of two exotic plants – Berberis thunbergii and Microstegium vimineum – in hardwood forests of New Jersey have shown a significant increase of pH in soils under the invasive plants as compared with soils from under native shrubs (Vaccinium spp). We present a further investigation of soil properties under the exotic plants in question. We measured the densities of earthworms in the soil under the two exotics and the native shrubs in three parks in New Jersey. In the same populations we also measured the extractable ammonium and nitrate in the top 5 cm of the soil, as well as the respiration of the soils and the potential rates of mineralization (aerobic lab incubation). In addition, we measured the nitrate reductase activity in leaves of the two exotic plants and several native shrubs and trees. Although there were differences between parks, we observed significantly higher earthworm densities in the soil under the exotic species. The worms were all European species. Soil pH, available nitrate and net potential nitrification were significantly higher in soils under the two exotic species. In contrast, total soil C and N and net ammonification were significantly higher under native vegetation. Nitrate reductase activities were much higher in the leaves of exotic plants than in the leaves of native shrubs and trees. Changes in soil properties, especially the change in nitrogen cycling, associated with the invasion of these two plant species may permit the invasion of other weedy or exotic species. Our results also suggest that even if the two exotic species were removed, the restoration of the native flora might be inhibited by the high nitrate concentrations in the soil.     

荒漠草原区柠条固沙人工林地表草本植被季节变化特征

研究荒漠草原人工林固沙区地表草本植被季节变化特征及其和柠条林龄的关系,对于分析柠条人工林地表草本植物的季节适应性和制订合理的人工林管理措施均具有重要的科学意义。选择6、15、24年生和36年生柠条人工林为研究对象,通过调查每个样地5月、8月和10月地表草本植物密度、物种数、盖度和高度,分析了荒漠草原区柠条人工固沙林生长过程中地表草本植被季节变化特征及其影响因素。结果表明,地表草本植物物种数在柠条林龄6和15a时受季节改变的影响较小,在24a之后受到季节变化的显著影响(P0.05)。地表草本植物密度在柠条林龄6a时受季节改变的影响较小,但在15a之后季节变化显著影响地表草本植物个体数分布(P0.05),而且在10月具有最多的地表草本植物个体数。地表草本植被盖度和高度均受到季节变化的显著影响(P0.05),而受林龄的影响较小;不同年龄林地地表草本植被盖度和高度均表现为10月和8月较高,5月较低。研究表明,荒漠草原柠条人工林固沙区,柠条林发育生长和灌木形态的改变不仅影响土壤营养条件,而且还可以调控由于季节改变而引起的土壤温湿度变化,柠条林龄和季节更替二者交互作用,共同影响地表草本植被的季节变化特征。     

Soil pH influences patterns of plant community composition after restoration with native‐based seed mixes

Reclamation of highly disturbed lands typically includes establishing fast‐growing, non‐native plants to achieve rapid ground cover for erosion control. Establishing native plant communities could achieve ecosystem functions beyond soil erosion, such as providing wildlife habitat. Pipelines, or other disturbed corridors through a landscape, present unique challenges for establishing native plant communities given the heterogeneity of soil environments and invasive plant propagule pressure. We created two structural equation models to address multiple related hypotheses about the influence of soil pH on plant community composition (current diversity and vegetative cover of the original restoration seed mix and background flora, and invasive plant density during mix establishment and current density) of a highly disturbed landscape corridor restored with native species. To test our hypotheses we conducted a plant survey on a gas pipeline crossing two state forests in the north‐central Appalachians that had been seeded with a native‐based mixture 8 years prior. Low soil pH was a strong predictor of density of the invasive annual plant, Microstegium vimineum, and had resulted in lower species diversity and cover of the seeded mix. Overall, our data provide evidence that native‐based grass and forb mixtures can establish and persist on a wide range of soil environments and thrive in competition with invasive plants in moderately acidic to neutral soils. Advancing knowledge on restoration methods using native species is essential to improving restoration practice norms to incorporate multifunctional ecological goals.     

Evaluation of a soil test method and plant analysis for determing the sulphur status of alluvial soils

Summary In a pot experiment with soils of Alfisol, Entisol, and Inceptisol orders, the relative yield of Egyptian clover (Trifolium alexandrinum L.) was significantly correlated with Morgan's reagent (N NaOAc+HOAc, pH 4.8)—extractable soil S (r=0.88), plant S (r=0.82), and plant N/S ratio (r=−0.77) suggesting suitability of these tests for diagnosing S deficiency. Total plant S lower than 0.21 per cent, plant N/S ratio wider than 17, and extractable soil S lower than 10 ppm were indicative of S deficiency, and were suggested therefore to be critical limits for these tests. Nitrogen and S in plant proteins were in near constant ratio of 16 and were significantly correlated (r=0.99). Sixty one per cent of 250 surface soil samples had less than 10 ppm extractable S and hence were deficient in S, suggesting a widespread S deficiency in soils under study. Extractable soil S in all soil series was significantly correlated with electrical conductivity and alkaline KMnO₄-extractable N, but not with pH, organic C, and CaCO₃.     

Release of Ni and Co by microbial activity in New Caledonian ultramafic soils

A high positive correlation was found between extractable Ni and Co contents and microbial activity of 40 ultramafic soil samples from New Caledonia, suggesting a possible role of microorganisms in the release of these two metals. A saprolite (ultramafic subsoil) and a hypermagnesian brown soil were incubated 9 months in different conditions. Ni and Co release, measured by their extractability, occurred without reduction of soil pH but did not occur when the native microflora was eliminated by heat treatment. However, when autoclaved soil was re-inoculated with a pinch of the same nonheated soil, the release of metals was recorded. The concentrations of extractable Ni and Co were much higher in soils amended with organic compounds in which microbial activity was enhanced, but only if the soil was not heat treated. The presence of Grevillea exul, a metal-tolerant plant, stimulated the metal release process, but the stimulation was less effective than it was in the compost-amended soil without plant. The influence of the plant in extractable Ni and Co contents in this amended soil was not significant. The release of the two metals therefore seemed to be induced by the activity of specialized organotrophic microorganisms.     

不同微生境下齿肋赤藓(Syntrichia caninervis)生理生化特性对不同季节的响应

齿肋赤藓(Syntrichia caninervis)作为典型的耐旱藓类,在古尔班通古特沙漠的藓类结皮中占优势地位。该沙漠的季节气候差异较大,冬季低温湿润,春季干旱,夏季高温且干旱。荒漠藓类植物叶片仅具单层细胞,对外界环境的变化十分敏感。而有关荒漠藓类植物在生理上如何适应这种剧烈环境变化还不得而知。研究测定了生长于两种不同微生境下的齿肋赤藓,经由低温湿润的冬季到干旱的春季再到高温干旱的夏季过程中生理生化变化特征,以探究不同微生境下齿肋赤藓在水热变化剧烈的不同季节的适应机制。研究发现:季节、微生境及二者的交互作用能够显著影响齿肋赤藓的游离脯氨酸、可溶性糖、可溶性蛋白及丙二醛(MDA)含量、过氧化氢酶(CAT)、过氧化物酶(POD)、超氧化物歧化酶(SOD)活性。夏季的高温干旱使齿肋赤藓的脯氨酸、可溶性糖、MDA含量及3种抗氧化酶活性均显著高于冬季及春季,而可溶性蛋白含量却呈相反趋势。干旱的春季齿肋赤藓脯氨酸及可溶性糖显著高于冬季。而在低温湿润的冬季,齿肋赤藓丙二醛含量及过氧化氢酶活性均显著高于春季。表明夏季齿肋赤藓所受胁迫最大,脯氨酸、可溶性糖含量及抗氧化酶活性大幅度提高。同时,在具有积雪覆盖的冬季,两种微生境下的齿肋赤藓生理生化特性无显著差异;而在无积雪覆盖的春夏季节,灌丛的遮阴作用为齿肋赤藓提供了水含量相对较高的良好生活环境,其下齿肋赤藓的渗透调节物质含量和抗氧化酶活性均显著低于裸露地。表明生长在裸露地的齿肋赤藓较活灌丛拥有更强的胁迫耐受性。