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• Background and Aims Differential responses of closely related species to submergence can provide insight into the evolution and mechanisms of submergence tolerance. Several traits of two wetland species from habitats with contrasting flooding regimes, Rorippa amphibia and Rorippa sylvestris, as well as F₁ hybrid Rorippa × anceps were analysed to unravel mechanisms underlying submergence tolerance. • Methods In the first submergence experiment (lasting 20 d) we analysed biomass, stem elongation and carbohydrate content. In the second submergence experiment (lasting 3 months) we analysed survival and the effect of re-establishment of air contact on biomass and carbohydrate content. In a separate experiment we analysed expression of two carbohydrate catabolism genes, ADH1 and SUS1, upon re-establishment of air contact following submergence. • Key Results All plants had low mortality even after 3 months of submergence. Rorippa sylvestris was characterized by 100 % survival and higher carbohydrate levels coupled with lower ADH1 gene expression as well as reduced growth compared with R. amphibia. Rorippa amphibia and the hybrid elongated their stems but this did not pay-off in higher survival when plants remained submerged. Only R. amphibia and the hybrid benefited in terms of increased biomass and carbohydrate accumulation upon re-establishing air contact. • Conclusions Results demonstrate contrasting 'escape' and 'quiescence' strategies between Rorippa species. Being a close relative of arabidopsis, Rorippa is an excellent model for future studies on the molecular mechanism(s) controlling these strategies.

Complete submergence represses photosynthesis and aerobic respiration, causing rapid mortality in most terrestrial plants. However, some plants have evolved traits allowing them to survive prolonged flooding, such as species of the genus Rorippa, close relatives of Arabidopsis (Arabidopsis thaliana). We studied plant survival, changes in carbohydrate and metabolite concentrations, and transcriptome responses to submergence of two species, Rorippa sylvestris and Rorippa amphibia. We exploited the close relationship between Rorippa species and the model species Arabidopsis by using Arabidopsis GeneChip microarrays for whole-genome transcript profiling of roots of young plants exposed to a 24-h submergence treatment or air. A probe mask was used based on hybridization of genomic DNA of both species to the arrays, so that weak probe signals due to Rorippa species/Arabidopsis mismatches were removed. Furthermore, we compared Rorippa species microarray results with those obtained for roots of submerged Arabidopsis plants. Both Rorippa species could tolerate deep submergence, with R. sylvestris surviving much longer than R. amphibia. Submergence resulted in the induction of genes involved in glycolysis and fermentation and the repression of many energy-consuming pathways, similar to the low-oxygen and submergence response of Arabidopsis and rice (Oryza sativa). The qualitative responses of both Rorippa species to submergence appeared roughly similar but differed quantitatively. Notably, glycolysis and fermentation genes and a gene encoding sucrose synthase were more strongly induced in the less tolerant R. amphibia than in R. sylvestris. A comparison with Arabidopsis microarray studies on submerged roots revealed some interesting differences and potential tolerance-related genes in Rorippa species.

Tetraploid inheritance has two extremes: disomic in allotetraploids and tetrasomic in autotetraploids. The possibility of mixed, or intermediate, inheritance models has generally been neglected. These could well apply to newly formed hybrids or to diploidizing (auto)tetraploids. We present a simple likelihood-based approach that is able to incorporate disomic, tetrasomic, and intermediate inheritance models and estimates the double-reduction rate. Our model shows that inheritance of microsatellite markers in natural tetraploids of Rorippa amphibia and R. sylvestris is tetrasomic, confirming their autotetraploid origin. However, in F1 hybrids inheritance was intermediate to disomic and tetrasomic inheritance. Apparently, in meiosis, chromosomes paired preferentially with the homolog from the same parental species, but not strictly so. Detected double-reduction rates were low. We tested the general applicability of our model, using published segregation data. In two cases, an intermediate inheritance model gave a better fit to the data than the tetrasomic model advocated by the authors. The existence of inheritance intermediate to disomic and tetrasomic has important implications for linkage mapping and population genetics and hence breeding programs of tetraploids. Methods that have been developed for either disomic or tetrasomic tetraploids may not be generally applicable, particularly in systems where hybridization is common.

Heterophylly, the phenomenon by which plants alter leaf forms to adapt to surrounding conditions, is apparent in amphibious plant species. In response to submergence, they emerge leaves with narrower blade areas. The pathway that receives the submergence signals and the mechanism regulating leaf form via cell proliferation and/or expansion systems have not yet been fully identified yet. Our anatomical study of Rorippa aquatica, an amphibious plant that exhibits heterophylly in response to various signals, showed that leaf thickness increased upon submergence; this was caused by the expansion of mesophyll cell size. Additionally, these submergence effects were inhibited under blue-light conditions. The ANGUSTIFOLIA3 (AN3)/GROWTH-REGULATING FACTOR (GRF) pathway regulating cell proliferation and cell expansion was downregulated in response to submergence; and the response was blocked under the blue-light conditions. These results suggest that submergence and light quality determine leaf cell morphology via the AN3/GRF pathway.

In order to maintain organs and structures at their appropriate sizes, multicellular organisms orchestrate cell proliferation and post-mitotic cell expansion during morphogenesis. Recent studies using Arabidopsis leaves have shown that compensation, which is defined as post-mitotic cell expansion induced by a decrease in the number of cells during lateral organ development, is one example of such orchestration. Some of the basic molecular mechanisms underlying compensation have been revealed by genetic and chimeric analyses. However, to date, compensation had been observed only in mutants, transgenics, and γ-ray-treated plants, and it was unclear whether it occurs in plants under natural conditions. Here, we illustrate that a shift in ambient temperature could induce compensation in Rorippa aquatica (Brassicaceae), a semi-aquatic plant found in North America. The results suggest that compensation is a universal phenomenon among angiosperms and that the mechanism underlying compensation is shared, in part, between Arabidopsis and R. aquatica.

Increasing the translocation and accumulation of cadmium (Cd) in Cd hyperaccumulator is an important technology to improve the phytoremediation efficiency of Cd-contaminated soil. In order to investigate the effects of different water conditions on the growth and Cd accumulation ability of Cd hyperaccumulators Rorippa sylvestris (L.) Besser and Rorippa amphibia Besser in Cd-polluted soil, clone seedlings of them were transplanted into pots filled with 50 mg kg −1 Cd-contaminated soil and cultured with water conditions of soil relative water content (RWC) 35%, 55%, 75%, 95%, and flooding respectively. The results showed the following: with the increase of RWC, the height of R. sylvestris and R. amphibia increased gradually, the dry biomass of shoot and whole plant increased and reached the maximum in 95% and then decreased in flooding; the Cd concentrations in shoots of R. sylvestris and R. amphibia were more than 100 mg kg −1 except for 35% and flooding; Cd bioconcentration factors (BCFs) of R. amphibia reached the maximum of 3.8870 in 75% and R. sylvestris reached the maximum of 3.2330 in 95%; sufficient water resulted in the decrease of photosynthetic rate due to more Cd accumulation. However, under flooding condition, because of the decrease of Cd bioavailability in soil, the accumulation of Cd in shoots declined and the net photosynthetic rate ( P n ) enhanced slightly.

Aim This article was aimed to explore the main rhizospherial properties of the Cd hyperaccumulator R. globosa compared to those of the non hyperaccumulator Rorippa palustris (Leyss.) Bess, representing the same genus (Rorippa) of Cruciferae. Method Pot culture experiments using soil spiked with Cd as CdCl₂ 2.5H₂O and rhizobags were conducted to determine the differences in Cd accumulation vs. pH, dissolved organic carbon (DOC), Cd chemical fractionation, enzyme activities, and microorganism number in the rhizospheres of R. globosa and R. palustris, and in the bulk soils. Results Experiments on Cd uptake by R. globosa and R. palustris from soil spiked with different doses of Cd ranging from 0 to 40 mgkg⁻¹, confirmed Cdhyperaccumulating properties of R. globosa (Cd accumulation in the above-ground organs > 100 mg kg⁻¹, enrichment factor EF >1, translocation factor TF >1, no significant biomass reduction at Cd doses > 10 mg kg⁻¹) and the lack of such properties in R. palustris, which made these species suitable for comparative studies. The pH value was found to be a constant, specific property of the rhizosphere of R. globosa and R. palustris, and of the bulk soil, independent on the Cd dose, however the differences were rather small: by 0.2 unit lower in the rhizosphere of R. globosa, and only by 0.1 unit lower in the rhizosphere of R.. palustris compared to the bulk soil. Chemical fractionation of Cd, i.e. its affinity to pools of different binding strength, also appeared to be a specific feature of a rhizosphere and soil independent on the Cd dose. It exhibited a unique capability of the rhizosphere of the Cd-hyperaccumulator R. globosa to mobilize Cd, which enriched the most labile exchangeable fraction in 24.4 % and the immobile residual fraction in 42.3 %, compared to 19.3 % and 50.8 % in the bulk soil and in the rhizosphere of the non-hiperaccumulator R. palustris that did not show significant difference (P< 0.05) from the bulk soil. In turn, DOC concentrations, enzymatic (urease and catalase) activity and microorganism (bacteria, fungi and actinomycetes) growth in rhizosphere soils were largely influenced by different Cd doses, although they were always considerably higher in the rhizosphere soils of R globosa, than in the rhizosphere oiR. palustris and in the bulk soil, in particular at Cd doses > 10 mg kg⁻¹. Conclusion pH and DOC changes in the rhizosphere of the Cd-hyperaccumulator R. globosa were found to be of a minor importance. The alteration of Cd chemical fractionation consisting in substantial reduction of the immobile residual pool and Cd enrichment primarily in the most labile exchangeable fraction, along with over 2-fold higher number of microorganisms was considered to be the driving force of Cd hyperaccumulation.

This experiment was used to explore whether the 11 nitrogenous nutrients affect the hyperaccumulation of Rorippa globosa (Turcz.) Thell. to Cd. Pot culture experiments using soil spiked with Cd as CdCl₂·2.5H₂O and 11 nitrogen-containing chemicals were conducted to determine the efficiency of the accumulation of Cd by R. globosa. Application of all 11 nitrogenous nutrients significantly (p < 0.05) enhanced Cd accumulation by R. globosa (Turcz.) Thell. Two major modes of Cd accumulation were observed: (i) through increase of biomass yield without reduction of Cd uptake and (ii) through increase of Cd uptake efficiency in parallel with increase of biomass yield. Bicarbonate > phosphate > chloride compounds of NH₄enhanced the biomass yield to the greatest extent, while oxalate > nitrate > chloride > and bicarbonate caused a significant increase of Cd uptake by R. globosa. Competition between N and Cd translocation caused either significant reduction of Cd translocation factor or decrease of biomass yield. Of studied nutrients, ammonium bicarbonate NH₄HCO₃and ammonium chloride NH₄Cl exerted the best joint effect of these two processes on the efficiency of R. globosa as a Cd hyperaccumulator. Application of these chemicals caused increase of Cd concentrations in roots of R. globosa by 35.1 and 41.1 %, and in shoots by 13.9 and 56.4 %, while biomasses of roots increased by 5.8- and 3.8-fold and in shoots by 7.4-fold, and 6.4-fold, respectively, compared to the control. As a result, accumulated load (μg pot⁻¹) of Cd in roots increased by 8.2- and 5.8-fold and in shoots by 8.6- and 10.6-fold in both pots. Consequently, chemicals (NH₄HCO₃and NH₄Cl) that enhanced both Cd enrichment and biomass yield had the greatest effect on the bioaccumulation capacity of R. globosa.

The river floodplain species Rorippa amphibia, Rorippa sylvestris, and their hybrid Rorippax anceps were studied here, with the aim of identifying potential species differences with respect to flooding tolerance, and of assessing their expression in F₁ hybrids. Parents and their F₁ hybrids were subjected to three flooding treatments mimicking natural conditions, and growth-related and leaf morphological traits were compared. In contrast to R. sylvestris, R. amphibia responded to waterlogging by forming specialized roots, and its growth was not reduced. These traits were dominantly expressed in hybrids. Both species and the hybrids established shoot growth over 2 wk of complete submergence. Only in R. sylvestris was this not at the expense of root biomass, suggesting that R. sylvestris can photosynthesize underwater. Rorippa sylvestris also showed a hyponastic response. Hybrids were intermediate to the parents in this respect. This study shows that phenotypic expression of parental traits in F₁ hybrids is mostly additive, but can also be dominant. This suggests that a large overlap in habitat use of parents and hybrids is likely. If such an overlap occurs, the main evolutionary consequences of hybridization in Rorippa will be the introgression of genes, as the hybrids are fully fertile.

The ability to respond to varying environments is crucial for sessile organisms such as plants. The amphibious plant Rorippa aquatica exhibits a striking type of phenotypic plasticity known as heterophylly, a phenomenon in which leaf form is altered in response to environmental factors. However, the underlying molecular mechanisms of heterophylly are yet to be fully understood. To uncover the genetic basis and analyze the evolutionary processes driving heterophylly in R. aquatica , we assembled the chromosome-level genome of the species. Comparative chromosome painting and chromosomal genomics revealed that allopolyploidization and subsequent post-polyploid descending dysploidy occurred during the speciation of R. aquatica . Based on the obtained genomic data, the transcriptome analyses revealed that ethylene signaling plays a central role in regulating heterophylly under submerged conditions, with blue light signaling acting as an attenuator of ethylene signal. The assembled R. aquatica reference genome provides insights into the molecular mechanisms and evolution of heterophylly. A chromosome-level genome assembly for the amphibious plant Rorippa aquatica uncovers its tetraploid origin and an involvement of ethylene in heterophylly upon submergence.