Evolution in stressful environments II: adaptive value and costs of plasticity in response to low light in Sinapis arvensis

Plants possess a remarkable capacity to alter their phenotype in response to the highly heterogeneous light conditions they commonly encounter in natural environments. In the present study with the weedy annual plant Sinapis arvensis, we (a) tested for the adaptive value of phenotypic plasticity in morphological and life history traits in response to low light and (b) explored possible fitness costs of plasticity. Replicates of 31 half-sib families were grown individually in the greenhouse under full light and under low light (40% of ambient) imposed by neutral shade cloth. Low light resulted in a large increase in hypocotyl length and specific leaf area (SLA), a reduction in juvenile biomass and a delayed onset of flowering. Phenotypic selection analysis within each light environment revealed that selection favoured large SLA under low light, but not under high light, suggesting that the observed increase in SLA was adaptive. In contrast, plasticity in the other traits measured was maladaptive (i.e. in the opposite direction to that favoured by selection in the low light environment). We detected significant additive genetic variance in plasticity in most phenotypic traits and in fitness (number of seeds). Using genotypic selection gradient analysis, we found that families with high plasticity in SLA had a lower fitness than families with low plasticity, when the effect of SLA on fitness was statistically kept constant. This indicates that plasticity in SLA incurred a direct fitness cost. However, a cost of plasticity was only expressed under low light, but not under high light. Thus, models on the evolution of phenotypic plasticity will need to incorporate plasticity costs that vary in magnitude depending on environmental conditions.     

Is seasonal variation in leaf traits adaptive for the annual plant Dicerandra linearifolia?

Empirical studies of phenotypic plasticity have often relied on the plausibility that a plastic response to the environment would increase fitness in order to diagnose the response as adaptive. I conducted a test of the hypothesis that seasonal variation in leaf traits is an adaptive response to seasonal variation in environmental conditions faced by the annual plant Dicerandralinearifolia. This species exhibits variation in leaf morphology and anatomy in response to temperature that is consistent with the expectations for adaptive plasticity. I examined variation in the size, thickness and density of stomata of leaves that develop in summer and winter and used analysis of phenotypic selection during winter and summer seasons to test the hypothesis that seasonal variation in these traits is adaptive. Regression analyses of estimated dry mass (as a proxy for fitness) on leaf traits revealed no evidence supporting the adaptive hypothesis. Selection favoured individuals with large and thick leaves in both winter and summer, and density of stomata had little or no effect on estimated relative fitness in any season. Correspondence between seasonal variation in leaf thickness and density of stomata and expectations for adaptive plasticity appears to be purely fortuitous. Seasonal variation in leaf traits may persist simply because there is no selection against individuals in which these traits vary. My results underscore the importance of definitive tests of the hypothesis of adaptation to distinguish adaptive plasticity from neutral or nonadaptive phenotypic plasticity.     

Selection on phenotypic plasticity of morphological traits in response to flooding and competition in the clonal shore plant Ranunculus reptans

Adaptive evolution of phenotypic plasticity requires that plastic genotypes have the highest global fitness. We studied selection by spatial heterogeneity of interspecific competition and flooding, and by temporal heterogeneity of flooding on morphological plasticity of 52 genotypes of the clonal shore plant Ranunculus reptans. Competition reduced clone size, rosette size, leaf length and stolon internode thickness. Flooding had similar effects and reduced competition. Differences in selection between environments imply potential for either local adaptation or for indirect evolution of phenotypic plasticity. We also detected direct selection for plastic reductions in internode length in response to flooding and for a plastic increase in internode length in response to competition. Plastic responses of some morphological traits to flooding were in line with selection thereon, suggesting that they indeed are adaptive and might have evolved in response to direct selection on plasticity.     

Phenotypic and transgenerational plasticity promote local adaptation to sun and shade environments

Adaptive plasticity is expected to be important when the grain of environmental variation is encompassed in offspring dispersal distance. We investigated patterns of local adaptation, selection and plasticity in an association of plant morphology with fine-scale habitat shifts from oak canopy understory to adjacent grassland habitat in Claytonia perfoliata. Populations from beneath the canopy of oak trees were >90 % broad leaved and large seeded, while plants from adjacent grassland habitat were >90 % linear-leaved and small seeded. In a 2-year study, we used reciprocal transplants and phenotypic selection analysis to investigate local adaptation, selection, plasticity and maternal effects in this trait-environment association. Transgenerational effects were studied by planting offspring of inbred maternal families grown in both environments across the same environments in the second year. Reciprocal transplants revealed local adaptation to habitat type: broad-leaved forms had higher fitness in oak understory and linear-leaved plants had higher fitness in open grassland habitat. Phenotypic selection analyses indicated selection for narrower leaves and lower SLA in open habitat, and selection for broad leaves and intermediate values of SLA in understory. Both plant morphs exhibited plastic responses in traits in the same direction as selection on traits (narrower leaves and lower SLA in open habitat) suggesting that plasticity is adaptive. We detected an adaptive transgenerational effect in which maternal environment influenced offspring fitness; offspring of grassland-reared plants had higher fitness than understory-reared plants when grown in grassland. We did not detect costs of plasticity, but did find a positive association between leaf shape plasticity and fitness in linear-leaved plants in grassland habitat. Together, these findings indicate that fixed differences in trait values corresponding to selection across habitat contribute to local adaptation, but that plasticity and maternal environmental effects may be favored through promotion of survival across heterogeneous environments.     

Differential expression of physiological and biochemical characters of some Indian mangroves towards salt tolerance

Mangroves are physiologically interesting as potential models for stress tolerance and as sources of alternative ideas about physiological strategies relevant at the ecosystem level. Variation in habitat has great impact on the physiological behavior and biochemical expression level of a particular plant species. Five species of mangroves, growing in saline and fresh water conditions were assessed for their ecological fitness in two different habitats. Assessments were based on some physiological and biochemical parameters measured from the fully exposed mature leaves under saline (15–27 PPT) and non-saline (1.2–2 PPT) conditions. Among the five species considered for investigation Bruguiera gymnorrhiza, Excoecaria agallocha and Phoenix paludosa grow luxuriously in the Sundarbans forest, while the rest two (Heritiera fomes, Xylocarpus granatum) are scanty. A comparative account of photosynthetic efficiency, chlorophyll content, mesophyll and stomatal conductance, specific leaf area, photosynthetic nitrogen use efficiency, total foliar free amino acids and differential expression of some antioxidant isoenzymes in leaf were estimated between the saline and non-saline plants. Elevated assimilation rate coupled with increased chlorophyll content, increased conductance and higher specific leaf area in non-saline condition indicates ability of these mangroves to grow even under minimal substrate salinity. The optimum PAR acquisition for photosynthesis in B. gymnorrhiza, E. agallocha and P. paludosa was higher under salt stress, while the maximum assimilation rate was lower in control plants. The opposite trend occurred in H. fomes and X. granatum, where the peak photosynthesis was lower under non-saline conditions even at a higher irradiance than in the saline forest. The isoform patterns of peroxidase, acid phosphatase and esterase indicated considerable difference in regulation of these enzymes due to salt stress and /or reverse adaptation.Key words: Chlorophyll content, Isozymes, Mangroves, Photosynthetic efficiency, Specific leaf area     

Testing the adaptive plasticity of Iris pumila leaf traits to natural light conditions using phenotypic selection analysis

A multivariate selection analysis has been used to test the adaptiveness of several Iris pumila leaf traits that display plasticity to natural light conditions. Siblings of a synthetic population comprising 31 families of two populations from contrasting light habitats were grown at an open dune site and in the understory of a Pinus nigra stand in order to score variation in phenotypic expression of six leaf traits: number of senescent leaves, number of live leaves, leaf length, leaf width, leaf angle, and specific leaf area. The ambient light conditions affected the values of all traits studied except for specific leaf area. In accordance to ecophysiological expectations for an adaptive response to light, both leaf length and width were significantly greater while the angle between sequential leaves was significantly smaller in the woodland understory than at the exposed dune site. The relationship between leaf traits and vegetative fitness (total leaf area) differed across light habitats as predicted by functional hypotheses. The standardized linear selection gradient (β′) for leaf length and width were positive in sign in both environments, but their magnitude for leaf length was higher in the shade than under full sunlight. Since plasticity of leaf length in the woodland shade has been recognized as adaptive, fitness cost of producing plastic change in leaf length was assessed. In both of the available methods used, the two-step and the multivariate regression procedures, a rather high negative association between the fitness value and the plasticity of leaf length was obtained, indicating a cost of plasticity. The selection gradient for leaf angle was weak and significant only in the woodland understory. Genetic correlations between trait expressions in contrasting light environments were negative in sign and low in magnitude, implying a significant genetic variation for plasticity in these leaf traits. Furthermore, leaf length and leaf width were found to be genetically positively coupled, which indicates that there is a potential for these two traits to evolve toward their optimal phenotypic values even faster than would be expected if they were genetically independent.     

Salinity Adaptation and the Contribution of Parental Environmental Effects in Medicago truncatula

High soil salinity negatively influences plant growth and yield. Some taxa have evolved mechanisms for avoiding or tolerating elevated soil salinity, which can be modulated by the environment experienced by parents or offspring. We tested the contribution of the parental and offspring environments on salinity adaptation and their potential underlying mechanisms. In a two-generation greenhouse experiment, we factorially manipulated salinity concentrations for genotypes of Medicago truncatula that were originally collected from natural populations that differed in soil salinity. To compare population level adaptation to soil salinity and to test the potential mechanisms involved we measured two aspects of plant performance, reproduction and vegetative biomass, and phenological and physiological traits associated with salinity avoidance and tolerance. Saline-origin populations had greater biomass and reproduction under saline conditions than non-saline populations, consistent with local adaptation to saline soils. Additionally, parental environmental exposure to salt increased this difference in performance. In terms of environmental effects on mechanisms of salinity adaptation, parental exposure to salt spurred phenological differences that facilitated salt avoidance, while offspring exposure to salt resulted in traits associated with greater salt tolerance. Non-saline origin populations expressed traits associated with greater growth in the absence of salt while, for saline adapted populations, the ability to maintain greater performance in saline environments was also associated with lower growth potential in the absence of salt. Plastic responses induced by parental and offspring environments in phenology, leaf traits, and gas exchange contribute to salinity adaptation in M. truncatula. The ability of plants to tolerate environmental stress, such as high soil salinity, is likely modulated by a combination of parental effects and within-generation phenotypic plasticity, which are likely to vary in populations from contrasting environments.     

Anatomical and physiological characteristics relating to ionic relations in some salt tolerant grasses from the Salt Range,Pakistan

Populations of three salt tolerant forage grasses (Cynodon dactylon, Imperata cylindrica, and Sporobolus arabicus) were collected from the salt-affected soils of the Salt Range and normal non-saline soils of the Faisalabad region to assess their mechanism of adaptation to saline stress by determining ion relations and some specific anatomical modifications. The population of S. arabicus from the Salt Range showed increased growth (root and shoot length, and root and shoot dry weights) under saline conditions. Salt tolerance in this species was related to structural modifications such as increased area of root, stem, leaf blade, and leaf sheath for toxic ion accumulation, increased vesicular hair density in leaves and aerenchyma formation in leaf sheath for ion exclusion. Uptake of toxic ions was high in the Salt Range population of C. dactylon and salt tolerance was related to ion exclusion through specific leaf structural modifications such as vesicular hairs. Salt tolerance in the Salt Range population of I. cylindrica was mainly associated with restricted uptake of toxic Na⁺ and Cl⁻ at root level, and accumulation of toxic ions via increased succulence in leaf blades and leaf sheaths in addition to some excretion of toxic ions through leaf sheath aerenchyma.     

Anatomical adaptations to salinity in cogon grass [Imperata cylindrica (L.) Raeuschel] from the Salt Range, Pakistan

To examine anatomical adaptations in a potential forage grass, Imperata cylindrica (L.) Raeuschel, a population was collected from the natural salt-affected soils of the Salt Range, Pakistan. Using a hydroponic system, the degree of salt tolerance in terms of structural modifications in the Salt Range ecotype was compared with that in an ecotype collected from a normal non-saline habitat of the Faisalabad region. The Salt Range ecotype was superior to the Faisalabad ecotype in biomass production under saline conditions. High salt tolerance of the Salt Range ecotype was associated with increased succulence in root and leaf (mainly midrib), formation of aerenchyma in leaf sheath, increased vascular bundle area, metaxylem area and phloem area, highly developed bulliform cells on leaves and increased sclerification in root and leaf. Furthermore, both stomatal density and stomatal area were considerably reduced under high salinities in the Salt Range ecotype.     

Adaptive plasticity in response to light and nutrient availability in the clonal plant Duchesnea indica

克隆植物蛇莓对光照强度和养分条件的适应性可塑性 表型可塑性可帮助植物缓冲环境压力并使其表型与当地环境相匹配,但目前仅少数性状的可塑性被广泛认为是适应性的。为充分理解可塑性的适应性意义,仍需进一步研究更多的植物功能性状及其环境因子。本研究将匍匐茎克隆植物蛇莓(Duchesnea indica)的21个基因型种植于不同的光照和养分条件下,并利用选择梯度分析检测了形态和生理可塑性对光照强度和养分有效性变化的适应性值。在遮荫条件下,蛇莓适合度(果实数、分株数和生物量)降低,节间缩短变细,成熟叶叶绿素含量降低,但叶柄长度、比叶面积、老叶叶绿素含量均增加。在低养分条件下,植株叶柄缩短,叶面积缩小变厚,叶绿素含量降低,但果实数量和根冠比增加。选择梯度分析表明,叶柄长度和老叶叶绿素含量对光照变化的可塑性是适应性的,老叶和成熟叶叶绿素含量对养分变化的可塑性也是适应性的。因此,不同性状的可塑性适应值取决于特定的生态背景。该研究的发现有助于理解克隆植物表型可塑性响应环境变化的适应性意义。     

The Invasion of Coastal Areas in South China by Ipomoea cairica May Be Accelerated by the Ecotype Being More Locally Adapted to Salt Stress

Local adaptation and phenotypic plasticity are two alternative mechanisms used by invasive plants for range expansion. We conducted a series of experiments to investigate the role of these mechanisms in the recent expansion of the invasive Ipomoea cairica from non-saline to salt-stressed coastal habitats. A comparison of the plant’s photosynthetic traits and construction costs across habitats was conducted through a field survey. Meanwhile, a full factorial greenhouse experiment was conducted with two ecotypes (non-saline and coastal) of I. cairica and two salinity gradients (water and 4 g L^-1 NaCl solution) to evaluate the roles of the two strategies by comparing their main traits. The results revealed that the construction cost and A_max of I. cairica did not change with the habitat type. The ecotype and saline treatments, however, significantly influenced the plant growth. The non-saline ecotype (NE) generally showed higher or equal plasticity of biomass-allocation and functional traits compared to the coastal ecotype (CE). However, the fitness and biomass of the NE significantly decreased with salinity, whereas those aspects of the CE did not change. Our results indicate that the recent expansion of I. cairica into coastal areas may be accelerated by the local adaptation of the CE to salt stress. Additionally, in South China, the CE will most likely evolve adaptations to both saline and non-saline environments, which will further broaden the invasion range of I. cairica in the future.     

旱地小麦理想株型研究进展

理想株型选择是旱地小麦节水栽培和高产育种的重要前提。围绕群体条件下旱地小麦产量形成、节水效率和干旱适应性等关键指标,从根型、茎型、叶型和穗型4个方面总结了旱地小麦株型演变规律和调控机理。旱地小麦从野生近缘种二倍体和四倍体向现代六倍体进化的漫长演变过程中,株型经历了从"高根冠比、低收获指数、高冠幅/株高比、小穗和种子扁长"等"杂草"型向"低根冠比、高收获指数和高粒叶比及小冠幅/株高、大穗多粒"的"作物"型的演变。从株型各组件演变规律看,根型以根系大小适中、根条数较多、根系生理活性较高为主;茎型以各茎节长度比例趋近"黄金分割"演变,株高控制在80-100 cm左右的适中高度;叶型以叶片直立、旗叶长宽比例适中为主,倒二叶长维持在20 cm左右;穗型以直立、大穗和小穗数多为主;单株分蘖数趋于减少,株型结构趋于紧凑。上述组件特征将促使旱地小麦向弱竞争能力的方向发展。总之,旱地小麦经历了自然选择和人工选择双重压力,其株型结构有利于群体产量和水分利用效率提高,从而获得较高的种群适合度。     

高寒草甸高原早熟禾个体性状对放牧与围封的响应

植物对放牧的响应是植物在响应过程中为生存和繁殖所形成的适应策略,研究植物表型反应对揭示草原生态系统的放牧响应机制具有重要意义。以青藏高原高寒草甸主要植物高原早熟禾(Poa alpigena)为研究对象,通过放牧和围封试验,研究其个体性状对放牧与围封的响应,旨在为草原生态系统放牧机制提供理论依据。结果表明:1)在长期放牧干扰下,除叶片数和根重外,其他性状株高、分枝数、叶长、叶宽、叶面积、茎粗、茎长、根长、根粗、穗长、总叶质量、单叶质量、茎重、穗重、全株重均出现显著变小的特征(P0.05);而短期围封对于高原早熟禾叶长、叶宽、总叶质量、单叶质量、茎重、穗重、根重等功能性状的恢复效果并不显著(P0.05),表明放牧退化草原植物性状具有保守性;2)通过构建高原早熟禾性状可塑性变化谱,发现穗重、茎重、全株重、分蘖数、茎长、株高等可塑性幅度较大,为放牧响应的敏感指标,叶片数、叶宽、茎粗、根粗的可塑性变化幅度较小,为惰性性状。     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Mechanisms driving plant functional trait variation in a tropical forest

Plant functional trait variation in tropical forests results from taxonomic differences in phylogeny and associated genetic differences, as well as, phenotypic plastic responses to the environment. Accounting for the underlying mechanisms driving plant functional trait variation is important for understanding the potential rate of change of ecosystems since trait acclimation via phenotypic plasticity is very fast compared to shifts in community composition and genetic adaptation. We here applied a statistical technique to decompose the relative roles of phenotypic plasticity, genetic adaptation, and phylogenetic constraints. We examined typically obtained plant functional traits, such as wood density, plant height, specific leaf area, leaf area, leaf thickness, leaf dry mass content, leaf nitrogen content, and leaf phosphorus content. We assumed that genetic differences in plant functional traits between species and genotypes increase with environmental heterogeneity and geographic distance, whereas trait variation due to plastic acclimation to the local environment is independent of spatial distance between sampling sites. Results suggest that most of the observed trait variation could not be explained by the measured environmental variables, thus indicating a limited potential to predict individual plant traits from commonly assessed parameters. However, we found a difference in the response of plant functional traits, such that leaf traits varied in response to canopy‐light regime and nutrient availability, whereas wood traits were related to topoedaphic factors and water availability. Our analysis furthermore revealed differences in the functional response of coexisting neotropical tree species, which suggests that endemic species with conservative ecological strategies might be especially prone to competitive exclusion under projected climate change.     

Evidence for plasticity,but not local adaptation,in invasive Japanese knotweed (<Emphasis Type="Italic">Reynoutria japonica</Emphasis>) in North America

While local adaptation and phenotypic plasticity are commonly observed in species occupying heterogeneous environments, these phenomena are less well understood in invasive species. However, plant invasions offer the opportunity to study these dynamics as they occur in species colonizing new habitats. In this study, we examined local adaptation and phenotypic plasticity in an invasive plant, Reynoutria japonica, across a broad latitudinal range within North America. We performed full-factorial reciprocal transplants using plants from three sites and examined fitness responses in both sexual and clonal reproductive measures, as well as vegetative responses in height, basal stem diameter, and biomass. For all vegetative traits, there was a significant effect of source population, indicating genetic differentiation among populations. There was also a significant effect of transplant site, suggesting phenotypic plasticity. However, there was no evidence of local adaptation at the North American meta-population level for either measure of sexual or clonal fitness. All three comparisons for sexual fitness failed to show any differences between source populations, indicating a lack of local adaptation. For clonal fitness, two of the three comparisons showed local maladaptation, and only one showed greater fitness at the home compared to foreign sites, but this population had greater fitness at all sites, indicating greater fitness overall for this population rather than local adaptation. The fact that we did not detect consistent patterns of local adaptation in these populations across a broad geographic range is somewhat surprising given that local adaptation appears common in many species, including invasives, and that the populations have been established for over a century. However, the lack of local adaptation observed in this species may indicate that phenotypic plasticity within the species is sufficient to allow the persistence of R. japonica in a variety of environments across its invaded range.     

Cytometric aspects of the leaves of the halophyte Suaeda maritima

An analysis of the frequency of occurrence of chloroplasts, mitochondria and microbodies in leaf cells of the halophyte Sueda maritima (L.) Dum, showed a remarkable increase (almost a doubling) in the number of mitochondria per cell section when plants were grown in saline (340 mol m⁻³ NaCl) as opposed to non-saline conditions. Chloroplast and microbody volume per cell increased under saline conditions, but number per cell was insensitive to salt. Chloroplasts contained a greater number of starch grains in plants grown under saline as compared to non-saline conditions and, under both conditions, were closely associated with the cell wall adjacent to the intercellular spaces. The implications for growth, photosynthesis and respiration are discussed.     

LOCAL ADAPTATION IN TWO SUBSPECIES OF AN ANNUAL PLANT: IMPLICATIONS FOR MIGRATION AND GENE FLOW

Plant populations often adapt to local environmental conditions. Here we demonstrate local adaptation in two subspecies of the California native annual Gilia capitata using standard reciprocal transplant techniques in two sites (coastal and inland) over three consecutive years. Subspecies performance in each site was measured in four ways: probability of seedling emergence, early vegetative size (length of longest leaf), probability of flowering, and total number of inflorescences produced per plant. Analysis of three of the four variables demonstrated local adaptation through site-by-subspecies interactions in which natives outperformed immigrants. The disparity between natives and immigrants in their probability of emergence and probability of flowering was greater at the coastal site than at the inland site. Treated in isolation, these two fitness components suggest that migration from the coast to the inland site may be less restricted by selection than migration in the opposite direction. Two measurements of individual size (leaf length and number of inflorescences), suggest (though not strongly) that immigrants may be subject to weaker selection at the coastal site than at the inland site. A standard cohort life table is used to compare replacement rates (R₀) for each subspecies at each site. Comparisons of R₀s suggest that immigrants are under a severe demographic disadvantage at the coastal site, but only a small disadvantage at the inland site. The results point out the importance of integrating over several fitness components when documenting the magnitude of local adaptation.     

Modulation of nutrient acquisition and polyamine pool in salt-stressed wheat (Triticum aestivum L.) plants inoculated with arbuscular mycorrhizal fungi

Two wheat (Triticum aestivum L.) cultivars, Sids 1 and Giza 168, were grown under non-saline or saline conditions (4.7 and 9.4 dS m^?1) with and without arbuscular mycorrhizal fungi (AMF) inoculation. Salt stress considerably decreased root colonization, plant productivity and N, P, K⁺, Fe, Zn and Cu concentrations, while it increased Na⁺ level, particularly in Giza 168. Mycorrhizal colonization significantly enhanced plant productivity and N, P, K⁺, Fe, Zn and Cu acquisition, while it diminished Na⁺ uptake, especially in Sids 1. Salinity increased putrescine level in Giza 168, however, values of spermidine and spermine increased in Sids 1 and decreased in Giza 168. Mycorrhization changed the polyamine balance under saline conditions, an increase in putrescine level associated with low contents of spermidine and spermine in Giza 168 was observed, while Sids 1 showed a decrease in putrescine and high increase in spermidine and spermine. Moreover, mycorrhizal inoculation significantly reduced the activities of diamine oxidase and polyamine oxidase in salt-stressed wheat plants. Modulation of nutrient acquisition and polyamine pool can be one of the mechanisms used by AMF to improve wheat adaptation to saline soils. This is the first report dealing with mycorrhization effect on diamine oxidase and polyamine oxidase activities under salt stress.     

Differential selection of growth rate-related traits in wild barley, Hordeum spontaneum, in contrasting greenhouse nutrient environments

Across-species comparisons show that inherent variation in relative growth rate (RGR) and its underlying traits are correlated with habitat productivity. In this study, we test the hypothesis that growth rate-related traits confer differential selective effects in contrasting nutrient environments. We specifically test whether high RGR is targeted by selection in nutrient-rich environments whereas low values of traits that underlie RGR [specific leaf area (SLA), leaf mass fraction and leaf area ratio (LAR)] confer a direct fitness advantage in nutrient-poor environments, resulting in selection of low RGR as a correlated response. We measured RGR, its underlying component traits, and estimated fitness in a range of wild barley (Hordeum spontaneum) accessions grown under high and low nutrient conditions. Selection on component traits differed between the two environments, while total selection of RGR was not significant. Using multiple regression and path analysis to estimate direct fitness effects, a selective advantage of high LAR and SLA was demonstrated only under nutrient-rich conditions. While supporting the view that observed associations between habitat richness and some RGR-component traits reflect adaptation to differing nutrient regimes, our data suggest that direct selection targets component traits rather than RGR itself.