Etiolated Seedling Development Requires Repression of Photomorphogenesis by a Small Cell-Wall-Derived Dark Signal.

Etiolated growth in darkness or the irreversible transition to photomorphogenesis in the light engages alternative developmental programs operating across all organs of a plant seedling. Dark-grown Arabidopsis de-etiolated by zinc (dez) mutants exhibit morphological, cellular, metabolic, and transcriptional characteristics of light-grown seedlings. We identify the causal mutation in TRICHOME BIREFRINGENCE encoding a putative acyl transferase. Pectin acetylation is decreased in dez, as previously found in the reduced wall acetylation2-3 mutant, shown here to phenocopy dez. Moreover, pectin of dez is excessively methylesterified. The addition of very short fragments of homogalacturonan, tri-galacturonate, and tetra-galacturonate, restores skotomorphogenesis in dark-grown dez and similar mutants, suggesting that the mutants are unable to generate these de-methylesterified pectin fragments. In combination with genetic data, we propose a model of spatiotemporally separated photoreceptive and signal-responsive cell types, which contain overlapping subsets of the regulatory network of light-dependent seedling development and communicate via a pectin-derived dark signal.

Relationships between soil and leaf mineral composition are element-specific, environment-dependent and geographically structured in the emerging model Arabidopsis halleri.

Leaf mineral composition, the leaf ionome, reflects the complex interaction between a plant and its environment including local soil composition, an influential factor that can limit species distribution and plant productivity. Here we addressed within-species variation in plant-soil interactions and edaphic adaptation using Arabidopsis halleri, a well-suited model species as a facultative metallophyte and metal hyperaccumulator. We conducted multi-element analysis of 1972 paired leaf and soil samples from 165 European populations of A. halleri, at individual resolution to accommodate soil heterogeneity. Results were further confirmed under standardized conditions upon cultivation of 105 field-collected genotypes on an artificially metal-contaminated soil in growth chamber experiments. Soil-independent between- and within-population variation set apart leaf accumulation of zinc, cadmium and lead from all other nutrient and nonessential elements, concurring with differential hypothesized ecological roles in either biotic interaction or nutrition. For these metals, soil-leaf relationships were element-specific, differed between metalliferous and nonmetalliferous soils and were geographically structured both in the field and under standardized growth conditions, implicating complex scenarios of recent ecological adaptation. Our study provides an example and a reference for future related work and will serve as a basis for the molecular-genetic dissection and ecological analysis of the observed phenotypic variation.

Is there a trade-off between glucosinolate-based organic and inorganic defences in a metal hyperaccumulator in the field?

Several plant species are able to not only tolerate but also hyperaccumulate heavy metals in their aboveground tissues. Thus, in addition to secondary metabolites acting as organic defences, metal hyperaccumulators possess an elemental defence that can act as protection against antagonists. Whereas several laboratory studies have determined potential relationships or trade-offs between organic and inorganic defences, little is known about whether these traits are interconnected in the field and which factors determine the compositions of organic defences and elements of leaf tissues most. To target these questions, we collected young leaves of Arabidopsis halleri, a Brassicaceae capable of hyperaccumulating Cd and Zn, as well as soil samples in the field from 16 populations. We detected wide variation in the composition of glucosinolates-the characteristic secondary metabolites of this plant family-among plants, with two distinct chemotypes occurring. Distance-based redundancy analyses revealed that variation in glucosinolate composition was determined mainly by population affiliation and to a lesser degree by geographic distance. Likewise, elemental composition of the leaves was mainly influenced by the location at which samples were collected. Therefore, the particular abiotic and biotic conditions and potential genetic relatedness at a particular locality affect the plant tissue chemistry. A slight indication of a trade-off between glucosinolate-based organic and inorganic defences was found, but only in the less abundant chemotype. A large variation in defence composition and potential joint effects of different defences may be highly adaptive ways of protecting against a wide arsenal of biotic antagonists.

Zinc and cadmium hyperaccumulation act as deterrents towards specialist herbivores and impede the performance of a generalist herbivore.

Extraordinarily high leaf metal concentrations in metal hyperaccumulator plants may serve as an elemental defence against herbivores. However, mixed results have been reported and studies using comparative approaches are missing. We investigated the deterrent and toxic potential of metals employing the hyperaccumulator Arabidopsis halleri. Effects of zinc (Zn) and cadmium (Cd) on the preferences of three Brassicaceae specialists were tested in paired-choice experiments using differently treated plant material, including transgenic plants. In performance tests, we determined the toxicity and joint effects of both metals incorporated in an artificial diet on the survival of a generalist. Feeding by all specialists was significantly reduced by metal concentrations from above 1000 µg Zn g(-1) DW and 18 µg Cd g(-1) DW. By contrast, metals did not affect oviposition. Generalist survival decreased with increasing concentrations of individual metals, whereby the combination of Zn and Cd had an additive toxic effect even at the lowest applied concentrations of 100 µg Zn g(-1) and 2 µg Cd g(-1) . Metal hyperaccumulation protects plants from herbivory resulting from deterrence and toxicity against a wide range of herbivores. The combination of metals exacerbates toxicity through joint effects and enhances elemental defence. Thus, metal hyperaccumulation is ecologically beneficial for plants.

Rosette iron deficiency transcript and microRNA profiling reveals links between copper and iron homeostasis in Arabidopsis thaliana.

Iron (Fe) is an essential plant micronutrient, and its deficiency limits plant growth and development on alkaline soils. Under Fe deficiency, plant responses include up-regulation of genes involved in Fe uptake from the soil. However, little is known about shoot responses to Fe deficiency. Using microarrays to probe gene expression in Kas-1 and Tsu-1 ecotypes of Arabidopsis thaliana, and comparison with existing Col-0 data, revealed conserved rosette gene expression responses to Fe deficiency. Fe-regulated genes included known metal homeostasis-related genes, and a number of genes of unknown function. Several genes responded to Fe deficiency in both roots and rosettes. Fe deficiency led to up-regulation of Cu,Zn superoxide dismutase (SOD) genes CSD1 and CSD2, and down-regulation of FeSOD genes FSD1 and FSD2. Eight microRNAs were found to respond to Fe deficiency. Three of these (miR397a, miR398a, and miR398b/c) are known to regulate transcripts of Cu-containing proteins, and were down-regulated by Fe deficiency, suggesting that they could be involved in plant adaptation to Fe limitation. Indeed, Fe deficiency led to accumulation of Cu in rosettes, prior to any detectable decrease in Fe concentration. ccs1 mutants that lack functional Cu,ZnSOD proteins were prone to greater oxidative stress under Fe deficiency, indicating that increased Cu concentration under Fe limitation has an important role in oxidative stress prevention. The present results show that Cu accumulation, microRNA regulation, and associated differential expression of Fe and CuSOD genes are coordinated responses to Fe limitation.

Use of natural variation reveals core genes in the transcriptome of iron-deficient Arabidopsis thaliana roots.

Iron (Fe) is an essential mineral micronutrient for plants and animals. Plants respond to Fe deficiency by increasing root uptake capacity. Identification of gene networks for Fe uptake and homeostasis could result in improved crop growth and nutritional value. Previous studies have used microarrays to identify a large number of genes regulated by Fe deficiency in roots of three Arabidopsis ecotypes. However, a large proportion of these genes may be involved in secondary or genotype-influenced responses rather than in a universal role in Fe uptake or homeostasis. Here we show that a small percentage of the Fe deficiency transcriptome of two contrasting ecotypes, Kas-1 and Tsu-1, was shared with other ecotypes. Kas-1 and Tsu-1 had different timing and magnitude of ferric reductase activity upon Fe withdrawal, and different categories of overrepresented Fe-regulated genes. To gain insights into universal responses of Arabidopsis to Fe deficiency, the Kas-1 and Tsu-1 transcriptomes were compared with those of Col-0, Ler, and C24. In early Fe deficiency (24-48 h), no Fe-downregulated genes and only 10 upregulated genes were found in all ecotypes, and only 20 Fe-downregulated and 58 upregulated genes were found in at least three of the five ecotypes. Supernode gene networks were constructed to visualize conserved Fe homeostasis responses. Contrasting gene expression highlighted different responses to Fe deficiency between ecotypes. This study demonstrates the use of natural variation to identify central Fe-deficiency-regulated genes in plants, and identified genes with potential new roles in signalling during Fe deficiency.