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Background and Aims Phenotypic plasticity allows plants to cope with environmental variability. Plastic responses to the environment have mostly been investigated at the level of individuals (plants) but can also occur within leaves. Yet the latter have been underexplored, as leaves are often treated as functional units with no spatial structure. We investigated the effect of a strong light gradient on plant and leaf traits and examined whether different portions of a leaf show similar or differential responses to light intensity. Methods We measured variation in 27 morpho-anatomical and physiological traits of the rosette and leaf portions (i.e. base and apex) of the tank bromeliad Aechmea aquilega (Bromeliaceae) when naturally exposed to a marked gradient of light intensity. Key Results The light intensity received by A. aquilega had a strong effect on the structural, biochemical and physiological traits of the entire rosette. Plants exposed to high light intensity were smaller and had wider, shorter, more rigid and more vertical leaves. They also had lower photosynthetic performance and nutrient levels. We found significant differences between the apex and basal portions of the leaf under low-light conditions, and the differences declined or disappeared for most of the traits as light intensity increased (i.e. leaf thickness, adaxial trichome density, abaxial and adaxial trichome surface, and vascular bundle surface and density). Conclusions Our results reveal a strong phenotypic plasticity in A. aquilega , particularly in the form of a steep functional gradient within the leaf under low-light conditions. Under high-light conditions, trait values were relatively uniform along the leaf. This study sheds interesting new light on the functional complexity of tank bromeliad leaves, and on the effect of environmental conditions on leaf trait regionalization. [ABSTRACT FROM AUTHOR]

Background and Aims Plants can respond to drought by changing their relative investments in the biomass and morphology of each organ. The aims of this study were to quantify the relative contribution of changes in morphology vs. allocation and determine how they affect each other. These results should help us understand the mechanisms that plants use to respond to drought events. Methods In a glasshouse experiment, we applied a drought treatment (well-watered vs. drought) at early and late stages of plant growth, leading to four treatment combinations (well-watered in both early and late periods, WW; drought in the early period and well-watered in the late period, DW; well-watered in the early period and drought in the late period, WD; drought in both early and late periods, DD). We used the variance partitioning method to compare the contribution of organ (leaf and root) biomass allocation and morphology to the leaf area ratio, root length ratio and root area ratio, for the rhizomatous grass Leymus chinensis (Trin.) Tzvelev. Key Results Compared with the continuously well-watered treatment, the leaf area ratio, root length ratio and root area ratio showed increasing trends under various drought treatments. The contribution of leaf mass allocation to leaf area ratio differed among the drought treatments and was 2.1- to 5.3-fold greater than leaf morphology, and the contribution of root mass allocation to root length ratio was ~2-fold greater than that of root morphology. In contrast, root morphology contributed more to the root area ratio than biomass allocation under drought in both the early and late periods. There was a negative correlation between the ratio of leaf mass fraction to root mass fraction and the ratio of specific leaf area to specific root length (or specific root area). Conclusions This study suggested that organ biomass allocation drove a larger proportion of variation than morphological traits for the absorption of resources in this rhizomatous grass. These findings should help us understand the adaptive mechanisms of plants when they are confronted with drought stress. [ABSTRACT FROM AUTHOR]

Background and Aims Cork oaks (Quercus section Cerris) comprise 15 extant species in Eurasia. Despite being a small clade, they display a range of leaf morphologies comparable to the largest sections (>100 spp.) in Quercus. Their fossil record extends back to the Eocene. Here, we explore how cork oaks achieved their modern ranges and how legacy effects might explain niche evolution in modern species of section Cerris and its sister section Ilex , the holly oaks. Methods We inferred a dated phylogeny for cork and holly oaks using a reduced-representation next-generation sequencing method, restriction site-associated DNA sequencing (RAD-seq), and used D- statistics to investigate gene flow hypotheses. We estimated divergence times using a fossilized birth–death model calibrated with 47 fossils. We used Köppen profiles, selected bioclimatic parameters and forest biomes occupied by modern species to infer ancestral climatic and biotic niches. Key Results East Asian and Western Eurasian cork oaks diverged initially in the Eocene. Subsequently, four Western Eurasian lineages (subsections) differentiated during the Oligocene and Miocene. Evolution of leaf size, form and texture was correlated, in part, with multiple transitions from ancestral humid temperate climates to mediterranean, arid and continental climates. Distantly related but ecologically similar species converged on similar leaf traits in the process. Conclusions Originating in temperate (frost-free) biomes, Eocene to Oligocene ranges of the primarily deciduous cork oaks were restricted to higher latitudes (Siberia to north of Paratethys). Members of the evergreen holly oaks (section Ilex) also originated in temperate biomes but migrated southwards and south-westwards into then-(sub)tropical southern China and south-eastern Tibet during the Eocene, then westwards along existing pre-Himalayan mountain ranges. Divergent biogeographical histories and deep-time phylogenetic legacies (in cold and drought tolerance, nutrient storage and fire resistance) thus account for the modern species mosaic of Western Eurasian oak communities, which are composed of oaks belonging to four sections. [ABSTRACT FROM AUTHOR]

Background and Aims Reproductive isolation and local establishment are necessary for plant speciation. Polyploidy, the possession of more than two complete chromosome sets, creates a strong postzygotic reproductive barrier between diploid and tetraploid cytotypes. However, this barrier weakens between polyploids (e.g. tetraploids and hexaploids). Reproductive isolation may be enhanced by cytotype morphological and environmental differentiation. Moreover, morphological adaptations to local conditions contribute to plant establishment. However, the relative contributions of ploidy level and the environment to morphology have generally been neglected. Thus, the extent of morphological variation driven by ploidy level and the environment was modelled for diploid, tetraploid and hexaploid cytotypes of Campanula rotundifolia agg. Cytotype distribution was updated, and morphological and environmental differentiation was tested in the presence and absence of natural contact zones. Methods Cytotype distribution was assessed from 231 localities in Central Europe, including 48 localities with known chromosome counts, using flow cytometry. Differentiation in environmental niche and morphology was tested for cytotype pairs using discriminant analyses. A structural equation model was used to explore the synergies between cytotype, environment and morphology. Key Results Tremendous discrepancies were revealed between the reported and detected cytotype distribution. Neither mixed-ploidy populations nor interploidy hybrids were detected in the contact zones. Diploids had the broadest environmental niche, while hexaploids had the smallest and specialized niche. Hexaploids and spatially isolated cytotype pairs differed morphologically, including allopatric tetraploids. While leaf and shoot morphology were influenced by environmental conditions and polyploidy, flower morphology depended exclusively on the cytotype. Conclusions Reproductive isolation mechanisms vary between cytotypes. While diploids and polyploids are isolated postzygotically, the environmental niche shift is essential between higher polyploids. The impact of polyploidy and the environment on plant morphology implies the adaptive potential of polyploids, while the exclusive relationship between flower morphology and cytotype highlights the role of polyploidy in reproductive isolation. [ABSTRACT FROM AUTHOR]

Background and Aims:Clines in traits across hybrid zones reflect a balance between natural selection and gene flow. Changes over time in average values for traits, and especially the shapes of their clines, are rarely investigated in plants, but could result from evolution in an unstable hybrid zone. Differences in clines between floral and vegetative traits could indicate different strengths of divergent selection. Methods:Five floral and two vegetative traits were measured in 12 populations along an elevational gradient spanning a natural hybrid zone between Ipomopsis aggregata and Ipomopsis tenuituba. We compared clines in the floral traits with those measured 25 years ago. Observed changes in mean trait values were compared with predictions based on prior estimates of natural selection. We also compared the steepness and position of clines between the floral and vegetative traits. Key Results:Corolla length has increased over five generations to an extent that matches predictions from measurements of phenotypic selection and heritability. The shape of its cline, and that of other traits, has not changed detectably. Clines varied across traits, but not all floral traits showed steeper clines than did vegetative traits. Both suites of morphological traits had steeper clines than did neutral molecular markers. Conclusions:The increase in corolla length provides a rare example of a match between predicted and observed evolution of a plant trait in natural populations. The clinal properties are consistent with the hypothesis that habitat-mediated divergent selection on vegetative traits and pollinator-mediated selection on floral traits both maintain species differences across the hybrid zone.

Background and Aims Leaf size has considerable ecological relevance, making it desirable to obtain leaf size estimations for as many species worldwide as possible. Current global databases, such as TRY, contain leaf size data for ~30 000 species, which is only ~8% of known species worldwide. Yet, taxonomic descriptions exist for the large majority of the remainder. Here we propose a simple method to exploit information on leaf length, width and shape from species descriptions to robustly estimate leaf areas, thus closing this considerable knowledge gap for this important plant functional trait. Methods Using a global dataset of all major leaf shapes measured on 3125 leaves from 780 taxa, we quantified scaling functions that estimate leaf size as a product of leaf length, width and a leaf shape-specific correction factor. We validated our method by comparing leaf size estimates with those obtained from image recognition software and compared our approach with the widely used correction factor of 2/3. Key Results Correction factors ranged from 0.39 for highly dissected, lobed leaves to 0.79 for oblate leaves. Leaf size estimation using leaf shape-specific correction factors was more accurate and precise than estimates obtained from the correction factor of 2/3. Conclusion Our method presents a tractable solution to accurately estimate leaf size when only information on leaf length, width and shape is available or when labour and time constraints prevent usage of image recognition software. We see promise in applying our method to data from species descriptions (including from fossils), databases, field work and on herbarium vouchers, especially when non-destructive in situ measurements are needed. [ABSTRACT FROM AUTHOR]

Background and Aims Intraspecific variation in foundation species of forest ecosystems can shape community and ecosystem properties, particularly when that variation has a genetic basis. Traits mediating interactions with other species are predicted by simple allocation models to follow ontogenetic patterns that are rarely studied in trees. The aim of this research was to identify the roles of genotype, ontogeny and genotypic trade-offs shaping growth, defence and reproduction in aspen. Methods We established a common garden replicating >500 aspen genets in Wisconsin, USA. Trees were measured through the juvenile period into the onset of reproduction, for growth, defence chemistry (phenolic glycosides and condensed tannins), nitrogen, extrafloral nectaries, leaf morphology (specific leaf area), flower production and foliar herbivory and disease. We also assayed the TOZ19 sex marker and heterozygosity at ten microsatellite loci. Key Results We found high levels of genotypic variation for all traits, and high heritabilities for both the traits and their ontogenetic trajectories. Ontogeny strongly shaped intraspecific variation, and trade-offs among growth, defence and reproduction supported some predictions while contradicting others. Both direct resistance (chemical defence) and indirect defence (extrafloral nectaries) declined during the juvenile stage, prior to the onset of reproduction. Reproduction was higher in trees that were larger, male and had higher individual heterozygosity. Growth was diminished by genotypic allocation to both direct and indirect defence as well as to reproduction, but we found no evidence of trade-offs between defence and reproduction. Conclusions Key traits affecting the ecological communities of aspen have high levels of genotypic variation and heritability, strong patterns of ontogeny and clear trade-offs among growth, defence and reproduction. The architecture of aspen's community genetics – its ontogeny, trade-offs and especially its great variability – is shaped by both its broad range and the diverse community of associates, and in turn further fosters that diversity. [ABSTRACT FROM AUTHOR]

Background and Aims Hybridization and introgression play an important role in the evolution and diversification of plants. To assess the degree of past and current hybridization, the level of genetic admixture in populations needs to be investigated. Ongoing hybridization and blurred species separation have made it challenging to assign European white oak taxa based on leaf morphology and/or genetic markers and to assess the level of admixture. Therefore, there is a need for powerful markers that differentiate between taxa. Here, we established a condensed set of single-nucleotide polymorphism (SNP) markers to reliably differentiate between the three most common oak species in temperate European forests (Quercus robur , Q. petraea , Q. pubescens) and to assess the degree of admixture in a large set of selected Swiss populations. Methods A training set of 194 presumably pure reference samples from Switzerland and Europe was used to assign 633 test individuals with two different approaches (population genetic-based/Bayesian vs. assumption-free/discriminative classifier) using 58 selected SNPs from coding regions. Admixture was calculated at the individual and population level with the Shannon diversity index based on individual assignment probabilities. Key Results Depending on the approach, 97.5–100 % of training individuals were assigned correctly, and additional analyses showed that the established SNP set could be further reduced while maintaining its discriminatory power. The two assignment approaches showed high overlap (99 %) in assigning training individuals and slightly less overlap in test individuals (84 %). Levels of admixture varied widely among populations. Mixed stands of Q. petraea and Q. pubescens revealed much higher degrees of admixture than mixed stands of the other two taxon pairs, accentuating high levels of gene flow between these two taxa in Switzerland. Conclusions Our set of SNPs warrants reliable taxon discrimination with great potential for further applications. We show that the three European white oak taxa have largely retained their species integrity in Switzerland despite high levels of admixture. [ABSTRACT FROM AUTHOR]

Background and Aims Differences in local abundance and ploidy level are predicted to impact the direction of introgression between species. Here, we tested these hypotheses on populations of Betula albosinensis (red birch) and Betula platyphylla (white birch) which were thought to differ in ploidy level, the former being tetraploid and the latter diploid. Methods We sampled 391 birch individuals from nine localities in China, and classified them into species based on leaf morphology. Twelve nuclear microsatellite markers were genotyped in each sample, and analysed using principal coordinates analysis and STRUCTURE software. We compared the effects of two different methods of scoring polyploid genotypes on population genetic analyses. We analysed the effect of ploidy, local species abundance and latitude on levels of introgression between the species. Key Results Leaf morphology divided our samples into red and white birch, but genetic analyses unexpectedly revealed two groups within red birch, one of which was tetraploid, as expected, but the other of which appeared to have diploid microsatellite genotypes. Five individuals were identified as early-generation hybrids or backcrosses between white birch and red birch and five were identified between red birch and 'diploid' red birch. Cline analysis showed that levels of admixture were not significantly correlated with latitude. Estimated genetic differentiation among species was not significantly different between determined tetraploid and undetermined tetraploid genotypes. Conclusions Limited hybridization and gene flow have occurred between red birch and white birch. Relative species abundance and ploidy level do not impact the direction of introgression between them, as genetic admixture is roughly symmetrical. We unexpectedly found populations of apparently diploid red birch and this taxon may be a progenitor of allotetraploid red birch populations. Incomplete lineage sorting may explain patterns of genetic admixture between apparently diploid and allotetraploid red birch. [ABSTRACT FROM AUTHOR]

Background and Aims Juvenile-to-adult phase transition is marked by changes in leaf morphology, mostly due to the temporal development of the shoot apical meristem, a phenomenon known as heteroblasty. Sugars and microRNA-controlled modules are components of the heteroblastic process in Arabidopsis thaliana leaves. However, our understanding about their roles during phase-changing in other species, such as Passiflora edulis , remains limited. Unlike Arabidopsis , P. edulis (a semi-woody perennial climbing vine) undergoes remarkable changes in leaf morphology throughout juvenile-to-adult transition. Nonetheless, the underlying molecular mechanisms are unknown. Methods Here we evaluated the molecular mechanisms underlying the heteroblastic process by analysing the temporal expression of microRNAs and targets in leaves as well as the leaf metabolome during P. edulis development. Key Results Metabolic profiling revealed a unique composition of metabolites associated with leaf heteroblasty. Increasing levels of glucose and α-trehalose were observed during juvenile-to-adult phase transition. Accumulation of microRNA156 (miR156) correlated with juvenile leaf traits, whilst miR172 transcript accumulation was associated with leaf adult traits. Importantly, glucose may mediate adult leaf characteristics during de novo shoot organogenesis by modulating miR156-targeted PeSPL9 expression levels at early stages of shoot development. Conclusions Altogether, our results suggest that specific sugars may act as co-regulators, along with two microRNAs, leading to leaf morphological modifications throughout juvenile-to-adult phase transition in P. edulis. [ABSTRACT FROM AUTHOR]

Background and Aims: Phenotypic plasticity allows plants to cope with environmental variability. Plastic responses to the environment have mostly been investigated at the level of individuals (plants) but can also occur within leaves. Yet the latter have been underexplored, as leaves are often treated as functional units with no spatial structure. We investigated the effect of a strong light gradient on plant and leaf traits and examined whether different portions of a leaf show similar or differential responses to light intensity., Methods: We measured variation in 27 morpho-anatomical and physiological traits of the rosette and leaf portions (i.e. base and apex) of the tank bromeliad Aechmea aquilega (Bromeliaceae) when naturally exposed to a marked gradient of light intensity., Key Results: The light intensity received by A. aquilega had a strong effect on the structural, biochemical and physiological traits of the entire rosette. Plants exposed to high light intensity were smaller and had wider, shorter, more rigid and more vertical leaves. They also had lower photosynthetic performance and nutrient levels. We found significant differences between the apex and basal portions of the leaf under low-light conditions, and the differences declined or disappeared for most of the traits as light intensity increased (i.e. leaf thickness, adaxial trichome density, abaxial and adaxial trichome surface, and vascular bundle surface and density)., Conclusions: Our results reveal a strong phenotypic plasticity in A. aquilega, particularly in the form of a steep functional gradient within the leaf under low-light conditions. Under high-light conditions, trait values were relatively uniform along the leaf. This study sheds interesting new light on the functional complexity of tank bromeliad leaves, and on the effect of environmental conditions on leaf trait regionalization., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Background and Aims In mature grass leaf blades as seen in cross-section, oblong cell-like structures have been interpreted most recently as intercellular gas spaces delimited by successive collapsed fusoid cells. These cells have been reported in at least seven of 12 subfamilies of Poaceae and are considered a synapomorphy for the family; however, no developmental work has been performed to verify their meristematic origin or to assess possible homologies within the graminid clade (= Flagellariaceae + [(Joinvilleaceae + Ecdeiocoleaceae) + Poaceae]) or among subfamilies of Poaceae. A developmental study was therefore carried out, including 20 species in three families (Flagellariaceae, Joinvilleaceae and Poaceae), representing the earlier-diverging and derived branches within the graminid clade and Poaceae. Methods Light microscopy was combined with scanning electron microscopy, cryoscanning electron microscopy and transmission electron microscopy to study the development of leaves taken from the shoot apex of young plants. Mature leaf blades also were taken from living or dried plants and the mid-portion was studied. Key Results Developmental results show that, in mature leaf blades as seen in cross-section, one apparent fusoid cell is typically a cavity resulting from the collapse of the initial fusoid cell and its internal divisions, which are herein interpreted as derivative cells with formation of cell plates only. Each cavity is delimited by successive collapsed fusoid cells arranged perpendicularly to the veins. Fusoid cells in all studied Poaceae members originate from the ground meristem, as do the colourless cells in Joinvillea ascendens (Joinvilleaceae). These two types of mesophyll cell have a strongly similar ontogeny, distinguished mainly by the collapse of the fusoid cells in Poaceae, which is not observed in the colourless cells in J. ascendens. Conclusions Within the Poaceae, the meristematic origin of fusoid cells is the same in the early-diverging lineages, BOP clade and Panicoideae, and thus they are homologous within the family. The same topography and meristematic origin suggest that fusoid cells in Poaceae and colourless cells in Joinvilleaceae are homologous. The results also suggest that the role played by the fusoid cells in young grass leaves is related to synthesis and storage of starch granules at early stages of development. [ABSTRACT FROM AUTHOR]

* Background and Aims The structural properties of leaf venation and xylem anatomy strongly influence leaf hydraulics, including the ability of leaves to maintain hydraulic function during drought. Here we examined the strength of the links between different leaf venation traits and leaf hydraulic vulnerability to drought (expressed as P50leaf by rehydration kinetics) in a diverse group of 26 woody angiosperm species, representing a wide range of leaf vulnerabilities, from four low-nutrient sites with contrasting rainfall across eastern Australia. v Methods For each species we measured key aspects of leaf venation design, xylem anatomy and leaf morphology. We also assessed for the first time the scaling relationships between hydraulically weighted vessel wall thickness (th) and lumen breadth (bh) across vein orders and habitats. * Key Results Across species, variation in P50leaf was strongly correlated with the ratio of vessel wall thickness (th) to lumen breadth (bh) [(t/b)h; an index of conduit reinforcement] at each leaf vein order. Concomitantly, the scaling relationship between th and bh was similar across vein orders, with a log-log slope less than 1 indicating greater xylem reinforcement in smaller vessels. In contrast, P50leaf was not related to th and bh individually, to major vein density (Dvmajor) or to leaf size. Principal components analysis revealed two largely orthogonal trait groupings linked to variation in leaf size and drought tolerance. * Conclusions Our results indicate that xylem conduit reinforcement occurs throughout leaf venation, and remains closely linked to leaf drought tolerance irrespective of leaf size. [ABSTRACT FROM AUTHOR]

Leaf anatomy, allometry, shape, size, leaf economics, leaf geometry Google Scholar Crossref Search ADS WorldCat 18 Niinmets U. 2015a. Is there a species spectrum within the world-wide leaf economics spectrum? Keywords: Leaf anatomy; allometry; shape; size; leaf geometry; leaf economics EN Leaf anatomy allometry shape size leaf geometry leaf economics i iii 1 07/12/23 20230509 NES 230509 Over the last two decades, trait-based plant ecology has been dominated by studies designed around the Leaf Economics Spectrum (LES), a framework that collapses variance in leaf traits onto a single axis based on the observed global trade-off between acquisitive and conservative investment strategies ([31]). [Extracted from the article]

Background and Aims Trait-environment relationships are commonly interpreted as evidence for local adaptation in plants. However, even when selection analyses support this interpretation, the mechanisms underlying differential benefits are often unknown. This study addresses this gap in knowledge using the broadly distributed South African shrub Protea repens. Specifically, the study examines whether broad-scale patterns of trait variation are consistent with spatial differences in selection and ecophysiology in the wild. Methods In a common garden study of plants sourced from 19 populations, associations were measured between five morphological traits and three axes describing source climates. Trait-trait and trait-environment associations were analysed in a multi-response model. Within two focal populations in the wild, selection and path analyses were used to test associations between traits, fecundity and physiological performance. Key Results Across 19 populations in a common garden, stomatal density increased with the source population's mean annual temperature and decreased with its average amount of rainfall in midsummer. Concordantly, selection analysis in two natural populations revealed positive selection on stomatal density at the hotter, drier site, while failing to detect selection at the cooler, moister site. Dry-site plants with high stomatal density also had higher stomatal conductances, cooler leaf temperatures and higher light-saturated photosynthetic rates than those with low stomatal density, but no such relationships were present among wet-site plants. Leaf area, stomatal pore index and specific leaf area in the garden also co-varied with climate, but within-population differences were not associated with fitness in either wild population. Conclusions The parallel patterns of broad-scale variation, differences in selection and differences in trait-ecophysiology relationships suggest a mechanism for adaptive differentiation in stomatal density. Densely packed stomata may improve performance by increasing transpiration and cooling, but predominately in drier, hotter climates. This study uniquely shows context-dependent benefits of stomatal density - a trait rarely linked to local adaptation in plants. [ABSTRACT FROM AUTHOR]

Background and Aims Leaf gas exchange is influenced by stomatal size, density, distribution between the leaf adaxial and abaxial sides, as well as by pore dimensions. This study aims to quantify which of these traits mainly underlie genetic differences in operating stomatal conductance (gs) and addresses possible links between anatomical traits and regulation of pore width.Methods Stomatal responsiveness to desiccation, gs-related anatomical traits of each leaf side and estimated gs (based on these traits) were determined for 54 introgression lines (ILs) generated by introgressing segments of Solanum pennelli into the S. lycopersicum ‘M82’. A quantitative trait locus (QTL) analysis for stomatal traits was also performed.Key Results A wide genetic variation in stomatal responsiveness to desiccation was observed, a large part of which was explained by stomatal length. Operating gs ranged over a factor of five between ILs. The pore area per stomatal area varied 8-fold among ILs (2–16 %), and was the main determinant of differences in operating gs between ILs. Operating gs was primarily positioned on the abaxial surface (60–83 %), due to higher abaxial stomatal density and, secondarily, to larger abaxial pore area. An analysis revealed 64 QTLs for stomatal traits in the ILs, most of which were in the direction of S. pennellii.Conclusions The data indicate that operating and maximum gs of non-stressed leaves maintained under stable conditions deviate considerably (by 45–91 %), because stomatal size inadequately reflects operating pore area (R2 = 0·46). Furthermore, it was found that variation between ILs in both stomatal sensitivity to desiccation and operating gs is associated with features of individual stoma. In contrast, genotypic variation in gs partitioning depends on the distribution of stomata between the leaf adaxial and abaxial epidermis. [ABSTRACT FROM PUBLISHER]

Background and Aims The study of intraspecific trait variation (ITV) in plants has a long history, dating back to the fourth century BC. Its existence was widely acknowledged by the end of the 18th century, although systematic and experimental studies commenced only a century later. However, the historiography of ITV has many gaps, especially with regard to early observations and visual documents. This note identifies an early depiction of plant ITV. Methods The botanical works of Johann Wolfgang von Goethe (1749–1832), a German writer and naturalist, were subjected to close reading. This included all publications and unpublished sources related to botany between 1785 and 1832 (e.g. notes, drafts, diaries, letters, drawings). This material is accessible in the multi-volume historical-critical edition of Goethe's studies in natural science (Leopoldina-Ausgabe). Key Results A diary entry from 9 September 1786 described changes in leaf morphology along an elevation gradient in the Bavarian Alps. The leaves of an unidentified species of willow (Spix sp.) and gentian (Gentiana sp.) were said to become narrower with increasing elevation; leaves also stood further apart on twigs, and the latter became thinner. A crude drawing of two willow twigs illustrated the differences. Goethe conjectured that the differences were due to environmental conditions. Conclusions Goethe's notes were anecdotal, and it is unclear whether the observed plant individuals actually belonged to the same species. Nevertheless, the notes represent an early and clear articulation of the hypothesis that changes in environmental conditions can cause ITV in a natural plant population. The drawing may be the earliest visual record of environmentally caused plant ITV in the wild. [ABSTRACT FROM AUTHOR]

Background and Aims A shift from outcrossing to selfing is thought to reduce the long-term survival of populations by decreasing the genetic variation necessary for adaptation to novel ecological conditions. However, theory also predicts an increase in adaptive potential as more of the existing variation becomes expressed as homozygous genotypes. So far, relatively few studies have examined how a transition to selfing simultaneously affects means, variances and covariances for characters that might be under stabilizing selection for a spatially varying optimum, e.g. characters describing leaf morphology. Methods Experimental crosses within an initially self-sterile population of Crepis tectorum were performed to produce an outbred and inbred progeny population to assess how a shift to selfing affects the adaptive potential for measures of leaf morphology, with special emphasis on the degree of leaf dissection, a major target of diversifying selection within the study species. Key Results Three consecutive generations of selfing had a minor impact on survival, the total number of heads produced and the mean leaf phenotype, but caused a proportional increase in the genetic (co)variance matrix for foliar characters. For the degree of leaf dissection, the lowest 50th percentile of the inbred progeny population showed a disproportionate increase in the genetic variance, consistent with the recessive nature of the weakly lobed phenotype observed in interpopulation crosses. Comparison of inbreeding response with large-scale patterns of variation indicates a potential for selection in a (recently) inbred population to drive a large evolutionary reduction in degree of leaf dissection by increasing the frequency of particular sibling lines. Conclusions The results point to a positive role for inbreeding in phenotypic evolution, at least during or immediately after a rapid shift in mating system. [ABSTRACT FROM AUTHOR]

Background and Aims Brassica rapa and B. oleracea are the progenitors of oilseed rape B. napus. The addition of each chromosome of B. oleracea to the chromosome complement of B. rapa results in a series of monosomic alien addition lines (MAALs). Analysis of MAALs determines which B. oleracea chromosomes carry genes controlling specific phenotypic traits, such as seed colour. Yellow-seeded oilseed rape is a desirable breeding goal both for food and livestock feed end-uses that relate to oil, protein and fibre contents. The aims of this study included developing a missing MAAL to complement an available series, for studies on seed colour control, chromosome homoeology and assignment of linkage groups to B. oleracea chromosomes. Methods A new batch of B. rapa–B. oleracea aneuploids was produced to generate the missing MAAL. Seed colour and other plant morphological features relevant to differentiation of MAALs were recorded. For chromosome characterization, Snow's carmine, fluorescence in situ hybridization (FISH) and genomic in situ hybridization (GISH) were used. Key Results The final MAAL was developed. Morphological traits that differentiated the MAALs comprised cotyledon number, leaf morphology, flower colour and seed colour. Seed colour was controlled by major genes on two B. oleracea chromosomes and minor genes on five other chromosomes of this species. Homoeologous pairing was largely between chromosomes with similar centromeric positions. FISH, GISH and a parallel microsatellite marker analysis defined the chromosomes in terms of their linkage groups. Conclusions A complete set of MAALs is now available for genetic, genomic, evolutionary and breeding perspectives. Defining chromosomes that carry specific genes, physical localization of DNA markers and access to established genetic linkage maps contribute to the integration of these approaches, manifested in the confirmed correspondence of linkage groups with specific chromosomes. Applications include marker-assisted selection and breeding for yellow seeds. [ABSTRACT FROM AUTHOR]

Background and Aims Plants have a family of proteins referred to as ICKs (inhibitors of cyclin-dependent kinase, CDK) or KRPs (Kip-related proteins) that function to regulate the activities of CDK. Knowledge of these plant CDK inhibitors has been gained mostly from studies of selected members in dicotyledonous plants, particularly Arabidopsis. Much remains to be learned regarding the differences among various members of the ICK/KRP family, and regarding the function and regulation of these proteins in monocotyledonous plants. Methods We analysed ICK-related sequences in the rice (Orysa sativa L. subsp. indica) genome and determined that there are six members with the conserved C-terminal signature region for ICK/KRP proteins. They are referred to as OsiICKs and further analyses were performed. The interactions with CDKs and cyclins were determined by a yeast two-hybrid assay, and cellular localization by fusion with the enhanced green fluorescence protein (EGFP). The expression of OsiICK6 in different tissues and in response to several treatments was analysed by reverse transcriptase-mediated polymerase chain reaction (RT-PCR) and real-time PCR. Furthermore, OsiICK6 was over-expressed in transgenic rice plants and significant phenotypes were observed. Key Results and Conclusions Based on putative protein sequences, the six OsiICKs are grouped into two classes, with OsiICK1 and OsiICK6 in each of the two classes, respectively. Results showed that OsiICK1 and OsiICK6 interacted with OsCYCD, but differed in their interactions with CDKA. Both EGFP:OsiICK1 and EGFP:OsiICK6 were localized in the nucleus. Whereas EGFP:OsiICK6 showed a punctuate subnuclear distribution, OsiICK1 had a homogeneous pattern. Over-expression of OsiICK6 resulted in multiple phenotypic effects on plant growth, morphology, pollen viability and seed setting. In OsiICK6-over-expressing plants, leaves rolled toward the abaxial side, suggesting that cell proliferation is critical in maintaining an even growth along the dorsal–ventral plane of leaf blades. [ABSTRACT FROM AUTHOR]

Background and Aims Understanding the fate and dynamics of cells during callus formation is essential to understanding totipotency and the mechanisms of somatic embryogenesis. Here, the fate of leaf explant cells during the development of embryogenic callus was investigated in the model legume Medicago truncatula. Methods Callus development was examined from cultured leaf explants of the highly regenerable genotype Jemalong 2HA (2HA) and from mesophyll protoplasts of 2HA and wild-type Jemalong. Callus development was studied by histology, manipulation of the culture system, detection of early production of reactive oxygen species and visualization of SERK1 (SOMATIC EMBRYO RECEPTOR KINASE1) gene expression. Key Results Callus formation in leaf explants initiates at the cut surface and within veins of the explant. The ontogeny of callus development is dominated by the division and differentiation of cells derived from pluripotent procambial cells and from dedifferentiated mesophyll cells. Procambium-derived cells differentiated into vascular tissue and rarely formed somatic embryos, whereas dedifferentiated mesophyll cells were competent to form somatic embryos. Interestingly, explants incubated adaxial-side down had substantially less cell proliferation associated with veins yet produced similar numbers of somatic embryos to explants incubated abaxial-side down. Somatic embryos mostly formed on the explant surface originally in contact with the medium, while in protoplast microcalli, somatic embryos only fully developed once at the surface of the callus. Mesophyll protoplasts of 2HA formed embryogenic callus while Jemalong mesophyll protoplasts produced callus rich in vasculature. Conclusions The ontogeny of embryogenic callus in M. truncatula relates to explant orientation and is driven by the dynamics of pluripotent procambial cells, which proliferate and differentiate into vasculature. The ontogeny is also related to de-differentiated mesophyll cells that acquire totipotency and form the majority of embryos. This contrasts with other species where totipotent embryo-forming initials mostly originate from procambial cells. [ABSTRACT FROM PUBLISHER]

Background and Aims Despite long-held interest, knowledge on why leaf size varies widely among species is still incomplete. This study was conducted to assess whether abiotic factors, phylogenetic histories and multi-trait interactions act together to shape leaf size. Methods Fifty-seven pairs of altitudinal vicariant species were selected in northern Spain, and leaf area and a number of functionally related leaf, shoot and whole plant traits were measured for each pair. Structural equation modelling helped unravel trait interactions affecting leaf size, and Mantel tests weighed the relative relevance of phylogeny, environment and trait interactions to explain leaf size reduction with altitude. Key Results Leaves of highland vicariants were generally smaller than those of lowlands. However, the extent of leaf size reduction with increasing altitude was widely variable among genera: from approx. 700 cm2 reduction (96 % in Polystichum) to approx. 30 cm2 increase (37 % in Sorbus). This was partially explained by shifts in leaf, shoot and whole plant traits (35–64 % of explained variance, depending on models), with size/number trade-offs more influential than shifts in leaf form and leaf economics. Shifts in traits were more important than phylogenetic distances or site-specific environmental variation in explaining the degree of leaf size reduction with altitude. Conclusions Ecological filters, constraints due to phylogenetic history (albeit modest in the study system), and phenotypic integration contribute jointly to shape single-trait evolution. Here, it was found that phenotypic change was far more important than shared ancestry to explaine leaf size differences of closely related species segregated along altitudes. [ABSTRACT FROM AUTHOR]

• Background Flooding causes substantial stress for terrestrial plants, particularly if the floodwater completely submerges the shoot. The main problems during submergence are shortage of oxygen due to the slow diffusion rates of gases in water, and depletion of carbohydrates, which is the substrate for respiration. These two factors together lead to loss of biomass and eventually death of the submerged plants. Although conditions under water are unfavourable with respect to light and carbon dioxide supply, photosynthesis may provide both oxygen and carbohydrates, resulting in continuation of aerobic respiration.• Scope This review focuses on evidence in the literature that photosynthesis contributes to survival of terrestrial plants during complete submergence. Furthermore, we discuss relevant morphological and physiological responses of the shoot of terrestrial plant species that enable the positive effects of light on underwater plant performance.• Conclusions Light increases the survival of terrestrial plants under water, indicating that photosynthesis commonly occurs under these submerged conditions. Such underwater photosynthesis increases both internal oxygen concentrations and carbohydrate contents, compared with plants submerged in the dark, and thereby alleviates the adverse effects of flooding. Additionally, several terrestrial species show high plasticity with respect to their leaf development. In a number of species, leaf morphology changes in response to submergence, probably to facilitate underwater gas exchange. Such increased gas exchange may result in higher assimilation rates, and lower carbon dioxide compensation points under water, which is particularly important at the low carbon dioxide concentrations observed in the field. As a result of higher internal carbon dioxide concentrations in submergence-acclimated plants, underwater photorespiration rates are expected to be lower than in non-acclimated plants. Furthermore, the regulatory mechanisms that induce the switch from terrestrial to submergence-acclimated leaves may be controlled by the same pathways as described for heterophyllous aquatic plants. [ABSTRACT FROM AUTHOR]

• Background and Aims Perennial plant formations always include a mixture of various-aged individuals of community-creating species, but the physiological and competitive potentials of plants of differing age and the importance on whole community functioning are still not entirely known. The current study tested the hypothesis that ontogenetically old plants have limited biomass investments in leaves and enhanced foliage support costs.• Methods Leaf structure, size and biomass allocation were studied in the perennial herb Pimpinella saxifraga during plant ontogeny from seedling to senile phases to determine age-dependent controls on key plant structural traits. The average duration of the full ontogenetic cycle is approx. 5–10 years in this species. Plants were sampled from shaded and open habitats.• Key Results Leaflet dry mass per unit area (MA) increased, and the fraction of plant biomass in leaflets (FL) decreased with increasing age, leading to a 5- to 11-fold decrease in leaf area ratio (LAR = FL/MA) between seedlings and senescent plants. In contrast, the fraction of below-ground biomass increased with increasing age. Leaflet size and number per leaf increased with increasing age. This was not associated with enhanced support cost in older plants as age-dependent changes in leaf shape and increased foliage packing along the rachis compensated for an overall increase in leaf size. Age-dependent trends were the same in habitats with various irradiance, but the LAR of plants of varying age was approx. 1·5-fold larger in the shade due to lower MA and larger FL.• Conclusions As plant light interception per unit total plant mass scales with LAR, these data demonstrate major age-dependent differences in plant light-harvesting efficiency that are further modified by site light availability. These ontogenetic changes reduce the differences among co-existing species in perennial communities, and therefore need consideration in our understanding of how herbaceous communities function. [ABSTRACT FROM AUTHOR]

• Background and Aims Trees with a partial leaf-shedding pattern and other morphological features a priori considered intermediate between those of the deciduous Nothofagus antarctica (G. Forster) Oersted and the evergreen N. dombeyi (Mirb.) Oersted (Nothofagaceae) were found in natural stands. The hybridization between a deciduous and an evergreen species of Nothofagus has not been reported so far in natural communities.• Methods The putative hybrids and the two presumed parental species were compared using 14 enzyme systems as well as shoot, leaf and reproductive morphology.• Key Results Six enzyme systems showed good resolution (MDH-B, IDH, SKDH, 6-PGDH, GOT and PGI) and in four of them (PGI, MDH-B, SKDH and 6-PGDH) the putative hybrids showed intermediate zymogram patterns between N. antarctica and N. dombeyi. Both principal coordinates analysis on isozyme data and principal components analysis (PCA) on quantitative morphological traits of shoots and leaves separated both parental species and located the putative hybrids closer to N. antarctica than to N. dombeyi. In the PCA, the number of basal cataphylls and the length : width ratio of leaves were the variables most discriminating among shoots of the three entities. The putative hybrids were intermediate between both species regarding leaf vernation, outline and venation, variation in leaf shape (length/width) with position on the parent shoot and in staminate inflorescence and cupule morphology. For other morphological traits, the putative hybrids resembled one of the parental species or differed from both species (e.g. valve morphology).• Conclusions Isoenzymatic and morphological data sets support the idea of the hybrid nature (probably F1 generation) of the semi-deciduous trees found. Nothofagus antarctica and N. dombeyi are probably more closely related than previously assumed. The relevance of pollen type in revealing evolutionary relationships between Nothofagus species is supported, and that of leaf-shedding pattern is rejected. [ABSTRACT FROM AUTHOR]

Leaf anatomy and morphology were studied in 11 tree species growing in an undisturbed forest and the adjoining fynbos for over 50 years. Functional anatomical results suggest that the forest and the fynbos are ecologically distinct. Moreover, leaf anatomy suggests that the foliage is primarily adapted for photosynthesis rather than for control of transpirational water loss. Forest precursor tree species and scrub species exhibit xeromorphy in the fynbos whereas they exhibit mesomorphic features inside the forest. The wide‐ranging species, such as Olea capensis subsp. capensis, simulated the response of the forest precursors, with the cuticle being phenotypically plastic between the forest and the fynbos but not between the stream and non‐stream habitats. Finally, the forest precursors, the scrub species, and the wide‐ranging taxa seem to have anatomical characters which can be modified in the fynbos and therefore allow its colonization by a variety of different species. [ABSTRACT FROM PUBLISHER]

A field experiment consisting of 18 birch trees grown in open top chambers in ambient and elevated CO2concentrations was set up with the aim of testing whether the positive growth response observed in many short-term studies is maintained after several growing seasons. We present the results of growth and biomass after 4.5 years of CO2exposure, one of the longest studies so far on deciduous tree species. We found that elevated CO2led to a 58% increase in biomass at the end of the experiment. However, estimation of stem mass during the growing season showed that elevated CO2did not affect relative growth rate during the fourth growing season, and therefore, that the large accumulation of biomass was the result of an early effect on relative growth rate in previous years. Trees grown in elevated CO2invested more carbon into fine roots and had relatively less leaf area than trees grown in ambient CO2. In contrast with previous studies, acceleration of growth did not involve a significant decline in nutrient concentrations of any plant tissue. It is likely that increased fine root density assisted the trees in meeting their nutrient demands. Changes in the species composition of the ectomycorrhizal fungi associated with the trees grown in elevated CO2in favour of late successional species supports the hypothesis of an acceleration of the ontogeny of the trees in elevated CO2.Copyright 1997 Annals of Botany Company [ABSTRACT FROM AUTHOR]

An introduction is presented in which the editor discusses articles in the issue on topics including ecology and significance of belowground bud banks, sex-biased gene expression in an annual dioecious herb, and ant pollination in Brazilian savanna.