Floristic characterisation of alkali soils in northwestern India

Summary Floristic composition studied in a 40 hectares representative site of alkali soils in the Indo-gangetic plains of Northwestern India revealed a remarkably restricted spectrum of natural vegetation. Soil analysis of specific niches, occupied by aggregations of different species, was employed to identify alkali halophytes. Response functions of eight prominent species to increasing levels of soil alkalinity, studied in pot culture, showed that most of them were adapted to alkali soils. Based on the computed importance value indices and observed biomass production maxima, it has been inferred thatSporobolus marginatus, Sporobulus coromandelianus, Diplachne fusca andChloris barbata, qualify as plant indicators of high-alkali soil conditions.     

Improved germination efficiency of Salicornia ramosissima seeds inoculated with Bacillus aryabhattai SP1016‐20

Saline agriculture and the crop cultivation of halophytes represent an alternative for the reclamation of salinized soils and for the management of irrigation water. Halotolerant plant growth promoting bacteria with biocontrol effect, as an alternative to commercial fungicides, may contribute to improve crop productivity while mitigating saline stress effects. The objective of this work was to isolate autochthonous rhizobacteria with biocontrol features, to be used as germination enhancers and plant‐growth promoting agents in the crop cultivation of Salicornia ramosissima. A set of isolates obtained from the rhizosphere of S. ramosissima was characterised in terms of Gram, motility, salt tolerance and biocontrol traits (hydrogen cyanide production, antifungal activity and production of extracellular lipases and proteases). One Gram‐positive motile isolate that tested positive for all biocontrol traits was identified by 16S rRNA gene sequencing as Bacillus aryabhattai. The inoculation of S. ramosissima seeds with B. aryabhattai SP1016‐20 reduced the negative effect of salinity on the germination efficiency. At the highest tested salinity (30 g/L NaCl) the final germination efficiency of inoculated seeds doubled in relation to non‐inoculated controls. Although the mechanisms involved in the biocontrol effect were not defined in the current work, the results highlight the potential of Bacillus aryabhattai SP1016‐20 as a plant‐growth promoting agent for the crop cultivation of Salicornia and contribute to the strengthening of the scientific basis of biosaline agriculture and plant growth promoting rhizobacteria‐assisted crop cultivation of halophytes in saline soils and estuarine sediments.     

Positive and negative biotic interactions and invasive Triadica sebifera tolerance to salinity: a cross‐continent comparative study

Exotic plant species may exhibit abiotic niche expansions that enable them to persist in a greater variety of habitat types in their introduced ranges than in their native ranges. This may reflect variation in limitation by different abiotic niche dimensions (realized niche shift) or phenotypic effects of biotic interactions that vary among ranges (realized niche expansion). Novel abiotic and biotic environments in the introduced range may also lead to genetic changes in exotic plant traits that enhance their abiotic stress tolerance (fundamental niche expansion). Here, we investigated how biotic interactions (aboveground herbivory and soil organisms) affect plant salinity tolerance using the invasive species Triadica sebifera from China (native range) and US (introduced range) populations grown in common gardens in both ranges. Simulated herbivory significantly reduced survival in saline treatments with reductions especially large at low salinity. Soil sterilization had a negative effect on survival at low salinity in China but had a positive effect on survival at low salinity in the US. Triadica survival and biomass were higher for US populations than for China populations, particularly in China but salinity tolerance did not depend on population origin. On average, arbuscular mycorrhizal (AM) colonization was higher for US populations, US soils and low salinity. These factors had a significant, positive, non‐additive interaction so that clipped seedlings from US populations in low saline US soils had high levels of AM colonization. Overall, our results show that phenotypic biotic interactions shape Triadica's salinity tolerance. Positive and negative biotic interactions together affected plant performance at intermediate stress levels. However, only aboveground damage consistently affected salinity tolerance, suggesting an important role for enemy release in expanding stress tolerance.     

Some studies on field distribution and seed germination ofSuaeda fruticosa andSporobolus arabicus with reference to salinity and sodicity of the medium

Summary Vegetation of two wasteland sites near Dera Chahl, 30 km from Lahore, was studied quantitatively. Soil samples obtained from the two sites were analysed.Suaeda fruticosa (L.) Forssk. was a dominant species on saline and sodic soil whereas soil underSporobolus arabicus Boiss. was saline and non-sodic. Seed germination studies show thatSporobolus arabicus is relatively more affected by Nasalinity than by Ca-salinity. This species tolerates salinity to some extent but is sensitive to sodicity.Suaeda fruticosa is relatively more tolerant to salinity and sodicity thanSporobolus arabicus and is capable of growing on saline and sodic soils.     

Phenotypic plasticity of four Chenopodiaceae species with contrasting saline–sodic tolerance in response to increased salinity–sodicity

It is unknown whether phenotypic plasticity in fitness‐related traits is associated with salinity–sodicity tolerance. This study compared growth and allocation phenotypic plasticity in two species with low salinity–sodicity tolerance (Chenopodium acuminatum and C. stenophyllum) and two species with high salinity–sodicity tolerance (Suaeda glauca and S. salsa) in a pot experiment in the Songnen grassland, China. While the species with low tolerance had higher growth and allocation plasticity than the highly tolerant species, the highly tolerant species only adjusted their growth traits and maintained higher fitness (e.g., plant height and total biomass) in response to increased soil salinity–sodicity, with low biomass allocation plasticity. Most plasticity is “apparent” plasticity (ontogenetic change), and only a few traits, for example, plant height:stem diameter ratio and root:shoot biomass ratio, represent “real” plasticity (real change in response to the environment). Our results show that phenotypic plasticity was negatively correlated with saline–sodic tolerance and could be used as an index of species sensitivity to soil salinity–sodicity.     

Enhanced growth of halophyte plants in biochar‐amended coastal soil: roles of nutrient availability and rhizosphere microbial modulation

Soil health is essential and irreplaceable for plant growth and global food production, which has been threatened by climate change and soil degradation. Degraded coastal soils are urgently required to reclaim using new sustainable technologies. Interest in applying biochar to improve soil health and promote crop yield has rapidly increased because of its multiple benefits. However, effects of biochar addition on the saline–sodic coastal soil health and halophyte growth were poorly understood. Response of two halophytes, Sesbania (Sesbania cannabina) and Seashore mallow (Kosteletzkya virginica), to the individual or co‐application of biochar and inorganic fertilizer into a coastal soil was investigated using a 52 d pot experiment. The biochar alone or co‐application stimulated the plant growth (germination, root development, and biomass), primarily attributed to the enhanced nutrient availability from the biochar‐improved soil health. Additionally, the promoted microbial activities and bacterial community shift towards the beneficial taxa (e.g. Pseudomonas and Bacillus) in the rhizosphere also contributed to the enhanced plant growth and biomass. Our findings showed the promising significance because biochar added at an optimal level (≤5%) could be a feasible option to reclaim the degraded coastal soil, enhance plant growth and production, and increase soil health and food security.     

Effects of salinity, external K+/Na+ ratio and soil moisture on growth and ion content of Sesbania rostrata

Growth of Sesbania rostrata was decreased gradually with increase in root medium salinity (mixed salts or NaCl alone). Soil moisture or anoxia did not affect plant growth significantly. Higher K⁺/Na⁺ ratios in plant tissues compared to those in the root medium were found under different salinities. This indicated a high K⁺-Na⁺ selectivity, a characteristic generally considered unique to halophytes. S. rostrata is moderately salt tolerant and may be utilized as forage crop and green manure on saline land. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diversity and salt tolerance of native Acacia rhizobia isolated from saline and non‐saline soils

Re‐establishing native vegetation in stressed soils is of considerable importance in many parts of the world, leading to significant interest in using plant–soil symbiont interactions to increase the cost‐effectiveness of large‐scale restoration. However, effective use of soil microbes in revegetation requires knowledge of how microbe communities vary along environmental stress gradients, as well as how such variation relates to symbiont effectiveness. In Australia, shrubby legumes dominate many ecosystems where dryland salinity is a major issue, and improving plant establishment in saline soils is a priority of regional management agencies. In this study, strains of rhizobial bacteria were isolated from a range of Acacia spp. growing in saline and non‐saline soils. Replicates of each strain were grown under several salinity levels in liquid culture and characterized for growth and salt tolerance. Genetic characterization of rhizobia showed considerable variation among strains, with salt tolerance and growth generally higher in rhizobial populations derived from more saline soils. These strains showed markedly different genetic profiles and generic affiliations to those from more temperate soils, suggesting community differentiation in relation to salt stress. The identification of novel genomic species from saline soils suggests that the diversity of rhizobia associated with Australian Acacia spp. is significantly greater than previously described. Overall, the ability of some symbiotically effective strains to tolerate high salinity is promising with regard to improving host plant re‐establishment in these soils.     

Naturally Saline Boreal Communities as Models for Reclamation of Saline Oil Sand Tailings

Reclaimed landscapes after oil sands mining have saline soils; yet, they are required to have similar biodiversity and productivity as the predisturbance nonsaline landscape. Given that many species in the boreal forest are not tolerant of salinity, we studied the effects of soil salinity on plant communities in natural saline landscapes to understand potential plant responses during the reclamation process. Vegetation–soil relationships were measured along transects from flooded wetlands to upland forest vegetation in strongly saline, slightly saline, nonsaline, and reclaimed boreal landscapes. In strongly saline landscapes, surface soil salinity was high (>10 dS/m) in flooded, wet‐meadow, and dry‐meadow vegetation zones as compared to slightly saline (<5 dS/m) and nonsaline (<2 dS/m) landscapes. Plant communities in these vegetation zones were quite different from nonsaline boreal landscapes and were dominated by halophytes common to saline habitats of the Great Plains. In the shrub and forest vegetation zones, surface soil salinity was similar between saline and nonsaline landscapes, resulting in similar plant communities. In strongly saline landscapes, soils remained saline at depth through the shrub and forest vegetation zones (>10 dS/m), suggesting that forest vegetation can establish over saline soils as long as the salts are below the rooting zone. The reclaimed landscape was intermediate between slightly saline and nonsaline landscapes in terms of soil salinity but more similar to nonsaline habitats with respect to species composition. Results from this study suggest it may be unrealistic to expect that plant communities similar to those found on the predisturbance landscape can be established on all reclaimed landscapes after oil sands mining.     

Biomass,anatomical changes and osmotic potential in Atriplex nummularia Lindl. cultivated in sodic saline soil under water stress

Atriplex nummularia exhibits excellent adaptability to environments with high salinity and low water availability. Accordingly, many studies have been conducted to identify the tolerance of the plant. We cultivated Atriplex in sodic saline soil under conditions of water stress in Northeast Brazil. The purpose of the study was to evaluate the growth characteristics and production of leaves, stems and roots of Atriplex under these conditions in order to identify anatomical changes in vesicular cells in leaf epidermis as well as to assess the osmotic potential of the soil solution and the leaves. The experiment was performed in a greenhouse where Atriplex was cultivated for 134 days in pots with sodic saline soil. The treatments comprised four moisture levels (35%, 55%, 75% and 95% of field capacity – FC). The height, diameter and dry mass of leaf, stem and root exhibited their highest values at levels of soil moisture that were 75% and 95% of FC. The high yields of dry biomass indicate the potential use of this halophyte for restoration of salt-affected soils. The vesicular cells were influenced by the soil moisture. The osmotic potential can serve as a good index for evaluating plant responses to water stress and salinity.     

Epidermal bladder cells confer salinity stress tolerance in the halophyte quinoa and Atriplex species

Epidermal bladder cells (EBCs) have been postulated to assist halophytes in coping with saline environments. However, little direct supporting evidence is available. Here, Chenopodium quinoa plants were grown under saline conditions for 5 weeks. One day prior to salinity treatment, EBCs from all leaves and petioles were gently removed by using a soft cosmetic brush and physiological, ionic and metabolic changes in brushed and non‐brushed leaves were compared. Gentle removal of EBC neither initiated wound metabolism nor affected the physiology and biochemistry of control‐grown plants but did have a pronounced effect on salt‐grown plants, resulting in a salt‐sensitive phenotype. Of 91 detected metabolites, more than half were significantly affected by salinity. Removal of EBC dramatically modified these metabolic changes, with the biggest differences reported for gamma‐aminobutyric acid (GABA), proline, sucrose and inositol, affecting ion transport across cellular membranes (as shown in electrophysiological experiments). This work provides the first direct evidence for a role of EBC in salt tolerance in halophytes and attributes this to (1) a key role of EBC as a salt dump for external sequestration of sodium; (2) improved K⁺ retention in leaf mesophyll and (3) EBC as a storage space for several metabolites known to modulate plant ionic relations.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.     

Effects of temperature,salinity and cold stratification on seed germination in halophytes

Salt tolerance of halophytes corresponds with the habitat requirement of the species. It is an important factor during the germination phase and it can determine successful establishment. This paper presents the effects of alternating temperature–light regimes (4/8°C, 10/20°C, 20/32°C; 12 h dark: 12 h light) and different salinity levels (0, 200, 400, 600 mmol l²¹ NaCl) on seed germination of five halophytes, Halimione pedunculata, Bupleurum tenuissimum, Aster tripolium, Triglochin maritimum and Armeria maritima. The five species differ with respect to family and life‐form and spatially correspond to a decreasing salt gradient (i.e. distance from salt water, with H. pedunculata being the most tolerant and A. maritima being the least). Armeria maritima, A. tripolium and T. maritimum seeds were additionally subjected to a cold stratification experiment. The results showed that Halimione pedunculata, an annual therophyte of year‐round heavily saline habitats, was dormant under all experimental conditions. Bupleurum tenuissimum, a species typical to sites of varying salinity prone to leaching during spring and autumn rainfall, germinated best under cold and warm temperatures, but only under non‐saline conditions. Aster tripolium and T. maritimum, close neighbours in salt marshes, showed very similar germination behaviour: seeds of both species tolerated high levels of salinity and germinated best in summer temperatures during periods of highest soil salinity, and germination was significantly promoted by cold. Armeria maritima, a species usually found on the marginal fringes of saline habitats, germinated only under low salt levels and maximum germination was under cold (spring) and warm (autumn) temperatures, with no significant effect of cold stratification.     

The effect of competition and edaphic conditions on the establishment of halophytes on brine effected soils

In order to test the feasibility of using native halophytes to reclaim brinecontaminated soil, seedlings of five inland halophytes, Atriplexprostrata, Hordeum jubatum, Salicornia europaea, Spergularia marina, and Suaeda calceoliformis, were planted at threedensities on a site near Athens, Ohio which had been contaminated by oilwell brine water. Ten replicates of each density treatment weretransplanted on two distinct areas of high and low salinity in May of 1993. Seedling survivorship, soil moisture, and soil salinity were monitored weeklythroughout the growing season. Biomass production and ion uptake weredetermined for each plant surviving until harvest. Soil analyses wereperformed prior to planting and after harvest to determine overall changesin soil chemistry and to determine the amount of Na⁺ reductionfrom the soil due to leaching by precipitation during the course of theexperiment. Survival was determined to be density independent for all ofthe species with the exception of S. marina where survival wasfacilitated at high density. Increased salinity negatively affected the survivaland yield of A. prostrata. The remaining species had greater survivalunder high salinity conditions, and density appeared to be the key factorinfluencing yield. Sodium and chloride ions were accumulated in planttissues in much greater amounts than K⁺, Ca⁺²or Mg⁺². Salicornia europaea plants grown in high densityon the high salinity site accumulated the highest amount of Na⁺ andH. jubatum grown in low density on the high salinity site accumulatedthe lowest amount of Na⁺. Soil salinities measured directly from theroot zone were significantly reduced (p<0.05) at the end of thegrowing season when compared to their controls. Atriplex prostrata(high density/low salinity) plots produced the greatest reduction in soilsalinity (15.8%) and S. marina (high density/high salinity) plots hadthe least reduction (1.2%).     

Effect of soil salinity and soil water availability on growth and chemical composition ofSorghum halepense L.

Summary Effects of soil salinity and soil water regime on growth and chemical composition ofSorghum halepense L. was studied with a view to evaluating its potential as a forage crop in saline soils. The experiment was conducted under controlled conditions using pot-culture with three levels of soil salinity (EC_e 0.5, 5.0, 10.0 ds/m) and three soil water regimes (60%, 40% and 20% of water holding capacity of the soil). High soil salinity and low soil water combiningly had an adverse effect on plant growth but the biomass production was appreciably high (57 to 75% of control) even under high soil salinity (EC_e 10 ds/m) when sufficient water was available. Belowground plant parts were relatively more salt-tolerant than shoots. There occurred an increase in the concentration of certain nutrients (N, Ca, Mg, TNC) in the plants in response to salinity, which along with increased root: shoot ratios was inferred as an adaptive feature of the plant for persistence under saline conditions.     

Morphological and structural plasticity of grassland species in response to a gradient in saline‐sodic soils

The abundance and distribution of species can be ascribed to both environmental heterogeneity and stress tolerance, with the latter measure sometimes associated with phenotypic plasticity. Although phenotypic plasticity varies predictably in response to common forms of stress, we lack a mechanistic understanding of the response of species to high saline‐sodic soils. We compared the phenotypic plasticity of three pairs of high and low saline‐sodic tolerant congeners from the families Poaceae (Leymus chinensis versus L. secalinus), Fabaceae (Lespedeza davurica versus L. bicolor) and Asteraceae (Artemisia mongolica versus A. sieversiana) in a controlled pot experiment in the Songnen grassland, China. The low tolerant species, L. secalinus and A. sieversiana exhibited higher plasticity in response to soil salinity and sodicity than their paired congeners. Highly tolerant species, L. chinensis and A. mongolica, had higher values for several important morphological traits, such as shoot length and total biomass under the high saline‐sodic soil treatment than their paired congeners. In contrast, congeners from the family Fabaceae, L. davurica and L. bicolor, did not exhibit significantly different plasticity in response to soil salinity and sodicity. All species held a constant reproductive effort in response to saline‐sodic soil stress. The different responses between low and high tolerant species offer an explanation for the distribution patterns of these species in the Songnen grassland. Highly tolerant species showed less morphological plasticity over a range of saline‐sodic conditions than their paired congeners, which may manifest as an inability to compete with co‐occurring species in locations where saline‐sodic soils are absent.     

MOVEMENT MODALITIES AND RESPONSES TO ENVIRONMENTAL CHANGES OF THE MUDFLAT DIATOM CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Cylindrotheca closterium (Ehrenberg) Reiman et Lewin is a raphid diatom widely distributed in mudflat assemblages. Video microscopy showed various movement modalities defined as smooth and corkscrew gliding, pirouette, pivot, rock and roll, rollover, and simultaneous pirouette and gliding. Z‐axis projection analysis of images revealed a unique gliding motif with corkscrew motions, which may have important ecological implications for C. closterium movement in muds. The general response to salinity alteration was a decrease in gliding movements with a concomitant increase in other modalities listed above. Short‐term responses to salinity change include dramatic alteration in modalities in hypo‐saline conditions and cessation of motility in extreme hyper‐saline environments. Modality changes were rapid and occurred within 5 s in response to hyper‐saline conditions. Hypo‐ or hyper‐saline conditions resulted in decreased gliding speed in standard media. Five‐ and 15‐day acclimation to salinity changes resulted in a progressive reduction in gliding movement, increased non‐gliding modalities and increased cell aggregation. Aggregation in hypo‐saline conditions was accompanied by a large increase in the polymer extracted by hot bicarbonate‐ and ethylenediamine tetraaceticacid‐ fractions of extracellular polymeric substance (EPS), the polymers of which have been implicated in cell attachment/motility phenomena. The monosaccharide profiles of these fractions were altered in response to hypo‐saline conditions. In general, monosaccharide profiles showed increased diversity upon cessation of motility and aggregation of cultures. The movement responses of C. closterium in response to environmental changes, accompanied by modifications in EPS, may form part of an adaptive strategy to survive in mudflats and could be useful as bioindicators of environmental changes.     

Reproductive allocation and fruit-set pattern in the euhalophyte Suaeda salsa in controlled and field conditions

Reproductive allocation and fruit-set pattern are important characteristics of halophytes, and these characteristics were investigated in euhalophyte Suaeda salsa (S. salsa). The plants comprised of those treated with 1 mM (control) or 500 mM (high salinity) of NaCl for 120 days in a glasshouse and those collected in the field. In controlled conditions, high salinity increased reproductive allocation and the number of fruits and seeds in each node of the branches. In field conditions, reproductive allocation and the number of fruits and brown seeds in S. salsa at the upper tide line were higher than at the lower tide line. There were more fruits and seeds distributed in the central part than those in the base or top of branches in both controlled and field conditions. Soil organic matter, total nitrogen and total phosphorus content at the upper tide line were lower than at the lower tide line. An opposite trend could be observed in the total soluble salt, Na⁺ and Cl^? content in soil. In conclusion, the fruit-set pattern of S. salsa may be affected by both resource constraints and resource allocation. Reproductive allocation and fruit-set pattern may ensure the establishment of S. salsa in high saline and low nutrient environments.     

Size advantage for male function and size‐dependent sex allocation in Ambrosia artemisiifolia,a wind‐pollinated plant

In wind‐pollinated plants, male‐biased sex allocation is often positively associated with plant size and height. However, effects of size (biomass or reproductive investment) and height were not separated in most previous studies. Here, using experimental populations of monoecious plants, Ambrosia altemisiifolia, we examined (1) how male and female reproductive investments (MRI and FRI) change with biomass and height, (2) how MRI and height affect male reproductive success (MRS) and pollen dispersal, and (3) how height affects seed production. Pollen dispersal kernel and selection gradients on MRS were estimated by 2,102 seeds using six microsatellite markers. First, MRI increased with height, but FRI did not, suggesting that sex allocation is more male‐biased with increasing plant height. On the other hand, both MRI and FRI increased with biomass but often more greatly for FRI, and consequently, sex allocation was often female‐biased with biomass. Second, MRS increased with both height and MRI, the latter having the same or larger effect on MRS. Estimated pollen dispersal kernel was fat‐tailed, with the maximum distance between mates tending to increase with MRI but not with height. Third, the number of seeds did not increase with height. Those findings showed that the male‐biased sex allocation in taller plants of A. artemisiifolia is explained by a direct effect of height on MRS.     

Relationships of reproductive performance indicators in black rhinoceros (Diceros bicornis michaeli) with plant available moisture,plant available nutrients and woody cover

Plant available moisture and plant available nutrients in soils influence forage quality and availability and subsequently affect reproductive performance in herbivores. However, the relationship of soil moisture, soil nutrients and woody forage with reproductive performance indicators is not well understood in mega‐browsers yet these three are important in selecting suitable areas for conservation of mega‐browsers. Here, the eastern black rhinoceros (Diceros bicornis michaeli), a mega‐browser, was studied in seven geographically distinct populations in Kenya to understand the relationships between its reproductive performance indicators and plant available moisture, plant available nutrients and woody cover. Reproductive parameters showed a complex relationship with plant available moisture and plant available nutrients. We found an increase in the predicted yearly percentage of females calving as plant available nutrients decreased in areas of high levels of plant available moisture but no relationship with plant available nutrients in areas of low plant available moisture. Age at first calving was earlier, inter‐calving interval was longer and yearly percentage of females calving was higher at higher woody cover. Woody plant cover contributes positively to black rhino reproduction performance indicators, whereas plant available moisture and plant available nutrients add to the selection of conservation areas, in more subtle ways.