Application of magnetic resonance imaging (MRI) technology in the characterization of microstructure and moisture content of young Moso bamboo.

Bamboo nodes play a key role in the hollow structure and the rapid growth of bamboo culm. Studying on the anatomical structure of bamboo is helpful to understand its growth mechanism. Taking the noninvasive, high-resolution and accurate technical advantages of magnetic resonance imaging (MRI), we conducted cross-sectional high-resolution MRI scanning on the tip of young Moso bamboo culm (removed shoot sheath) and extracted the gray value of the MRIs by using MATLAB software to explore the differences of water distribution in nodes, proximal nodes, and internodes. The results showed that numerous vascular bundles were repeatedly twisted and rotated horizontally at the nodal diaphragms and inner wall near the nodal diaphragms of the young bamboo, forming an intricate and highly connected network. The structure protected important tissues from mechanical stress by allocating axial loads, and enabled to laterally transport water and nutrients, which was an important basis for the rapid growth of Moso bamboo in relatively short term. The signal value (also known as brightness value) of MRIs indicated that water content of vascular bundles in young bamboo culm was much higher than that of surrounding parenchyma tissues. The mean value and standard deviation of water content between pixels of internodes were significantly higher than that of nodes, and the values of that in the proximal nodes were intermediate. The development of MRI would play a significant role in the studies of bamboo anatomy, physiology, and biochemistry.

If self-shading is so bad, why is there so much? Short shoots reconcile costs and benefits.

If trees minimize self-shading, new foliage in shaded parts of the crown should remain minimal. However, many species have abundant foliage on short shoots inside their crown. In this paper, we test the hypothesis that short shoots allow trees to densify their foliage in self-shaded parts of the crown thanks to reduced costs. Using 30 woody species in Mediterranean and tropical biomes, we estimated the contribution of short shoots to total plant foliage, calculated their costs relative to long shoots including wood cost and used 3D plant simulations calibrated with field measurements to quantify their light interception, self-shading and yield. In species with short shoots, leaves on short shoots account for the majority of leaf area. The reduced cost of short stems enables the production of leaf area with 36% less biomass. Simulations show that although short shoots are more self-shaded, they benefit the plant because they cost less. Lastly, the morphological properties of short shoots have major implications for whole plant architecture. Taken together, our results question the validity of only assessing leaf costs to understand leaf economics and call for more integrated observations at the crown scale to understand light capture strategies in woody plants.

Nitrogen Uptake and Distribution in Different Chinese Cabbage Genotypes under Low Nitrogen Stress.

In order to understand the effects of low nitrogen (LN) stress on the growth and development in different genotypes of Chinese cabbage, the L40 genotype with high nitrogen utilization and the L14 genotype with LN utilization were selected as experimental materials. Field experiments and indoor hydroponic methods were used to study the different responses of two Chinese cabbage genotypes to low nitrogen levels. In this study, we also analyzed the genome-wide gene expression profiles of L40 and L14 in response to LN stress by high-throughput RNA sequencing technology. The results reveal that the L40 root system responds better to LN compared with L14. After LN stress, L40 can effectively absorb and transport NO3- and store it in the ground. It is precisely because of this characteristic of the L40 genotype that LN treatment did not have a significant effect on the chlorophyll (Chl) content and net photosynthetic rate (Pn) of the L40 Chinese cabbage compared with the L14 Chinese cabbage. These two different Chinese cabbage genotypes were shown to have differently expressed genes related to nitrate transport, auxin synthesis, and glutamate dehydrogenase synthesis. These genes function in the nitrogen pathway, which are important candidates for understanding the molecular host-response mechanisms to LN stress.

Plasticity of bud outgrowth varies at cauline and rosette nodes in Arabidopsis thaliana.

Shoot branching is a complex mechanism in which secondary shoots grow from buds that are initiated from meristems established in leaf axils. The model plant Arabidopsis (Arabidopsis thaliana) has a rosette leaf growth pattern in the vegetative stage. After flowering initiation, the main stem elongates with the top leaf primordia developing into cauline leaves. Meristems in Arabidopsis initiate in the axils of rosette or cauline leaves, giving rise to rosette or cauline buds, respectively. Plasticity in the process of shoot branching is regulated by resource and nutrient availability as well as by plant hormones. However, few studies have attempted to test whether cauline and rosette branching are subject to the same plasticity. Here, we addressed this question by phenotyping cauline and rosette branching in three Arabidopsis ecotypes and several Arabidopsis mutants with varied shoot architectures. Our results showed no negative correlation between cauline and rosette branch numbers in Arabidopsis, demonstrating that there is no tradeoff between cauline and rosette bud outgrowth. Through investigation of the altered branching pattern of flowering pathway mutants and Arabidopsis ecotypes grown in various photoperiods and light regimes, we further elucidated that the number of cauline branches is closely related to flowering time. The number of rosette branches has an enormous plasticity compared with cauline branches and is influenced by genetic background, flowering time, light intensity, and temperature. Our data reveal different levels of plasticity in the regulation of branching at rosette and cauline nodes, and promote a framework for future branching analyses.

Distinct responses of frond and root to increasing nutrient availability in a floating clonal plant.

Current knowledge on responses of aquatic clonal plants to resource availability is largely based on studies manipulating limited resource levels, which may have failed to capture the "big picture" for aquatic clonal plants in response to resource availability. In a greenhouse experiment, we grew the floating clonal plant Spirodela polyrhiza under ten nutrient levels (i.e., 1/64×, 1/32×, 1/16×, 1/8×, 1/4×, 1/2×, 1×, 2×, 4× and 8×full-strength Hoagland solution) and examined their responses in terms of clonal growth, morphology and biomass allocations. The responses of total biomass and number of ramets to nutrient availability were unimodal. A similar pattern was found for frond mass, frond length and frond width, even though area per frond and specific frond area fluctuated greatly in response to nutrient availability. In contrast, the responses of root mass and root length to nutrient availability were U-shaped. Moreover, S. polyrhiza invested more to roots under lower nutrient concentrations. These results suggest that nutrient availability may have distinct influences on roots and fronds of the aquatic clonal plant S. polyrhiza, resulting in a great influence on the whole S. polyrhiza population.

Soil fertility interactions with Sinorhizobium-legume symbiosis in a simulated Martian regolith; effects on nitrogen content and plant health.

Due to increasing population growth and declining arable land on Earth, astroagriculture will be vital to terraform Martian regolith for settlement. Nodulating plants and their N-fixing symbionts may play a role in increasing Martian soil fertility. On Earth, clover (Melilotus officinalis) forms a symbiotic relationship with the N-fixing bacteria Sinorhizobium meliloti; clover has been previously grown in simulated regolith yet without bacterial inoculation. In this study, we inoculated clover with S. meliloti grown in potting soil and regolith to test the hypothesis that plants grown in regolith can form the same symbiotic associations as in soils and to determine if greater plant biomass occurs in the presence of S. meliloti regardless of growth media. We also examined soil NH4 concentrations to evaluate soil augmentation properties of nodulating plants and symbionts. Greater biomass occurred in inoculated compared to uninoculated groups; the inoculated average biomass in potting mix and regolith (2.23 and 0.29 g, respectively) was greater than the uninoculated group (0.11 and 0.01 g, respectively). However, no significant differences existed in NH4 composition between potting mix and regolith simulant. Linear regression analysis results showed that: i) symbiotic plant-bacteria relationships differed between regolith and potting mix, with plant biomass positively correlated to regolith-bacteria interactions; and, ii) NH4 production was limited to plant uptake yet the relationships in regolith and potting mix were similar. It is promising that plant-legume symbiosis is a possibility for Martian soil colonization.

What shoots can teach about theories of plant form.

Plants generate a large variety of shoot forms with regular geometries. These forms emerge primarily from the activity of a stem cell niche at the shoot tip. Recent efforts have established a theoretical framework of form emergence at the shoot tip, which has empowered the use of modelling in conjunction with biological approaches to begin to disentangle the biochemical and physical mechanisms controlling form development at the shoot tip. Here, we discuss how these advances get us closer to identifying the construction principles of plant shoot tips. Considering the current limits of our knowledge, we propose a roadmap for developing a general theory of form development at the shoot tip.

Arabidopsis SMAX1 overaccumulation suppresses rosette shoot branching and promotes leaf and petiole elongation.

ARABIDOPSIS: SMAX1/SMXL (SUPPRESSOR OF MAX2 1/SMAX1-LIKE) proteins function as transcriptional repressors in karrikin and strigolactone (SL) signaling pathways and regulate plant architecture. MAX2 is a common factor in the two signaling pathways and a component of the SCF complex that modulates the proteasome-mediated degradation of SMAX1/SMXLs. SMXL6, 7, and 8 proteins promote shoot branching and inhibit petiole elongation. Our study found that the accumulation of SMAX1 suppresses rosette shoot branching and increases cauline branches on the primary inflorescence stem, plant height, petiole length, and leaf length/width ratio. The SMAX1 accumulation enhances the expression of BRC1, HB53, HB40, and HB21 that modulate shoot branching. SMAX1 also regulates the expression of the genes involved in auxin transport, cytokinin signaling pathway, and SL biosynthesis. The expression analyses of these genes suggest that excessive SMAX1 should accelerate the transport of auxin and the biosynthesis of SL in plants. High SL concentration suppresses the bud development in smax1D mutant that accumulates SMAX1 protein in plant. However, the effects of cytokinin and auxin on shoot branching remain elusive in the mutant with excessive SMAX1. SMAX1 regulates leaf shape and petiole length via modulating TCP1 expression. Our findings reveal a novel function of SMAX1 and new mechanism of shoot branching.

Insights on Molecular Characteristics of Hydrochars by 13C-NMR and Off-Line TMAH-GC/MS and Assessment of Their Potential Use as Plant Growth Promoters.

Hydrochar is a carbon-based material that can be used as soil amendment. Since the physical-chemical properties of hydrochar are mainly assigned to process parameters, we aimed at evaluating the organic fraction of different hydrochars through 13C-NMR and off-line TMAH-GC/MS. Four hydrochars produced with sugarcane bagasse, vinasse and sulfuric or phosphoric acids were analyzed to elucidate the main molecular features. Germination and initial growth of maize seedlings were assessed using hydrochar water-soluble fraction to evaluate their potential use as growth promoters. The hydrochars prepared with phosphoric acid showed larger amounts of bioavailable lignin-derived structures. Although no differences were shown about the percentage of maize seeds germination, the hydrochar produced with phosphoric acid promoted a better seedling growth. For this sample, the greatest relative percentage of benzene derivatives and phenolic compounds were associated to hormone-like effects, responsible for stimulating shoot and root elongation. The reactions parameters proved to be determinant for the organic composition of hydrochar, exerting a strict influence on molecular features and plant growth response.

Potential transceptor AtNRT1.13 modulates shoot architecture and flowering time in a nitrate-dependent manner.

Compared with root development regulated by external nutrients, less is known about how internal nutrients are monitored to control plasticity of shoot development. In this study, we characterize an Arabidopsis thaliana transceptor, NRT1.13 (NPF4.4), of the NRT1/PTR/NPF family. Different from most NRT1 transporters, NRT1.13 does not have the conserved proline residue between transmembrane domains 10 and 11; an essential residue for nitrate transport activity in CHL1/NRT1.1/NPF6.3. As expected, when expressed in oocytes, NRT1.13 showed no nitrate transport activity. However, when Ser 487 at the corresponding position was converted back to proline, NRT1.13 S487P regained nitrate uptake activity, suggesting that wild-type NRT1.13 cannot transport nitrate but can bind it. Subcellular localization and ß-glucuronidase reporter analyses indicated that NRT1.13 is a plasma membrane protein expressed at the parenchyma cells next to xylem in the petioles and the stem nodes. When plants were grown with a normal concentration of nitrate, nrt1.13 showed no severe growth phenotype. However, when grown under low-nitrate conditions, nrt1.13 showed delayed flowering, increased node number, retarded branch outgrowth, and reduced lateral nitrate allocation to nodes. Our results suggest that NRT1.13 is required for low-nitrate acclimation and that internal nitrate is monitored near the xylem by NRT1.13 to regulate shoot architecture and flowering time.

Interactions between two QTLs for time to anthesis on spike development and fertility in wheat.

Earliness per se (Eps) genes are reported to be important in fine-tuning flowering time in wheat independently of photoperiod (Ppd) and vernalisation (Vrn). Unlike Ppd and Vrn genes, Eps have relatively small effects and their physiological effect along with chromosomal position are not well defined. We evaluated eight lines derived from crossing two vernalisation insensitive lines, Paragon and Baj (late and early flowering respectively), to study the detailed effects of two newly identified QTLs, Eps-7D and Eps-2B and their interactions under field conditions. The effect of both QTLs was minor and was affected by the allelic status of the other. While the magnitude of effect of these QTLs on anthesis was similar, they are associated with very different profiles of pre-anthesis development which also depends on their interaction. Eps-7D affected both duration before and after terminal spikelet while not affecting final leaf number (FLN) so Eps-7D-early had a faster rate of leaf appearance. Eps-2B acted more specifically in the early reproductive phase and slightly altered FLN without affecting the leaf appearance rate. Both QTLs affected the spike fertility by altering the rate of floret development and mortality. The effect of Eps-2B was very small but consistent in that -late allele tended to produce more fertile florets.

Environmental and genetic regulation of plant height in soybean.

BACKGROUND: Shoot architecture is fundamentally crucial to crop growth and productivity. As a key component of shoot architecture, plant height is known to be controlled by both genetic and environmental factors, though specific details remain scarce. RESULTS: In this study, 308 representative soybean lines from a core collection and 168 F9 soybean progeny were planted at distinct field sites. The results demonstrated the presence of significant genotype × environment interaction (G × E) effects on traits associated with plant height in a natural soybean population. In total, 19 loci containing 51 QTLs (quantitative trait locus) for plant height were identified across four environments, with 23, 13 and 15 being QTLs for SH (shoot height), SNN (stem node number) and AIL (average internode length), respectively. Significant LOD ranging from 2.50 to 16.46 explained 2.80-26.10% of phenotypic variation. Intriguingly, only two loci, Loc11 and Loc19-1, containing 20 QTLs, were simultaneously detected across all environments. Results from Pearson correlation analysis and PCA (principal component analysis) revealed that each of the five agro-meteorological factors and four soil properties significantly affected soybean plant height traits, and that the corresponding QTLs had additive effects. Among significant environmental factors, AD (average day-length), AMaT (average maximum temperature), pH, and AN (available nitrogen) had the largest impacts on soybean plant height. Therefore, in spite of uncontrollable agro-meteorological factors, soybean shoot architecture might be remolded through combined efforts to produce superior soybean genetic materials while also optimizing soil properties. CONCLUSIONS: Overall, the comprehensive set of relationships outlined herein among environment factors, soybean genotypes and QTLs in effects on plant height opens new avenues to explore in work aiming to increase soybean yield through improvements in shoot architecture.

The Potassium Transporter OsHAK5 Alters Rice Architecture via ATP-Dependent Transmembrane Auxin Fluxes.

Plant HAK/KUP/KT family members function as plasma membrane (PM) H+/K+ symporters and may modulate chemiosmotically-driven polar auxin transport (PAT). Here, we show that inactivation of OsHAK5, a rice K+ transporter gene, decreased rootward and shootward PAT, tiller number, and the length of both lateral roots and root hairs, while OsHAK5 overexpression increased PAT, tiller number, and root hair length, irrespective of the K+ supply. Inhibitors of ATP-binding-cassette type-B transporters, NPA and BUM, abolished the OsHAK5-overexpression effect on PAT. The mechanistic basis of these changes included the OsHAK5-mediated decrease of transmembrane potential (depolarization), increase of extracellular pH, and increase of PM-ATPase activity. These findings highlight the dual roles of OsHAK5 in altering cellular chemiosmotic gradients (generated continuously by PM H+-ATPase) and regulating ATP-dependent auxin transport. Both functions may underlie the prominent effect of OsHAK5 on rice architecture, which may be exploited in the future to increase crop yield via genetic manipulations.

OsHOX1 and OsHOX28 Redundantly Shape Rice Tiller Angle by Reducing HSFA2D Expression and Auxin Content.

Tiller angle largely determines plant architecture, which in turn substantially influences crop production by affecting planting density. A recent study revealed that HEAT STRESS TRANSCRIPTION FACTOR2D (HSFA2D) acts upstream of LAZY1 (LA1) to regulate tiller angle establishment in rice (Oryza sativa). However, the mechanisms underlying transcriptional regulation of HSFA2D remain unknown. In this study, two class II homeodomain-Leu zipper genes, OsHOX1 and OsHOX28, were identified as positive regulators of tiller angle by affecting shoot gravitropism. OsHOX1 and OsHOX28 showed strong transcriptional suppressive activity in rice protoplasts and formed intricate self- and mutual-transcriptional negative feedback loops. Moreover, OsHOX1 and OsHOX28 bound to the pseudopalindromic sequence CAAT(C/G)ATTG within the promoter of HSFA2D, thus suppressing its expression. In contrast to HSFA2D and LA1, OsHOX1 and OsHOX28 attenuated lateral auxin transport, thus repressing the expression of WUSCHEL-RELATED HOMEOBOX 6 (WOX6) and WOX11 in the lower side of the shoot base of plants subjected to gravistimulation. Genetic analysis further confirmed that OsHOX1 and OsHOX28 act upstream of HSFA2D Additionally, both OsHOX1 and OsHOX28 inhibit the expression of multiple OsYUCCA genes and decrease auxin biosynthesis. Taken together, these results demonstrated that OsHOX1 and OsHOX28 regulate the local distribution of auxin, and thus tiller angle establishment, through suppression of the HSFA2D-LA1 pathway and reduction of endogenous auxin content. Our finding increases the knowledge concerning fine tuning of tiller angles to optimize plant architecture in rice.

Life in the Current: Anatomy and Morphology of Utricularia neottioides.

Rheophytism is extremely rare in the Utricularia genus (there are four strictly rheophytic species out of a total of about 260). Utricularia neottioides is an aquatic rheophytic species exclusively growing attached to bedrocks in the South American streams. Utricularia neottioides was considered to be trap-free by some authors, suggesting that it had given up carnivory due to its specific habitat. Our aim was to compare the anatomy of rheophytic U. neottioides with an aquatic Utricularia species with a typical linear monomorphic shoot from the section Utricularia, U. reflexa, which grows in standing or very slowly streaming African waters. Additionally, we compared the immunodetection of cell wall components of both species. Light microscopy, histochemistry, scanning, and transmission electron microscopy were used to address our aims. In U. neottioides, two organ systems can be distinguished: organs (stolons, inflorescence stalk) which possess sclerenchyma and are thus resistant to water currents, and organs without sclerenchyma (leaf-like shoots), which are submissive to the water streaming/movement. Due to life in the turbulent habitat, U. neottioides evolved specific characters including an anchor system with stolons, which have asymmetric structures, sclerenchyma and they form adhesive trichomes on the ventral side. This anchor stolon system performs additional multiple functions including photosynthesis, nutrient storage, vegetative reproduction. In contrast with typical aquatic Utricularia species from the section Utricularia growing in standing waters, U. neottioides stems have a well-developed sclerenchyma system lacking large gas spaces. Plants produce numerous traps, so they should still be treated as a fully carnivorous plant.

Use of Zinc Carbonate Spiking to Obtain Phytotoxicity Thresholds Comparable to Those in Field-Collected Soils.

Several studies have reported the presence of smithsonite (ZnCO3 ) in soils polluted by zinc mining. The present study aimed to determine upper critical threshold values of Zn phytotoxicity in a substrate spiked with ZnCO3 and to compare them with those obtained in field-collected soils. We studied Zn toxicity to perennial ryegrass (Lolium perenne L.) grown in pots with unpolluted peat treated with increasing concentrations of ZnCO3 that produced nominal total Zn concentrations of 0, 0.7, 1.3, 2.0, 2.6, and 3.3%. To keep constant near-neutral pH value in all the treatments, we used decreasing concentrations of dolomitic lime. In the treatment with total soil Zn of 3.3% (pH 6.8), the foliar Zn concentration of L. perenne was 1914 ± 211 mg kg-1 , falling into the range of 2400 ± 300 mg kg-1 reported for Lolium species grown under similar laboratory conditions in a polluted soil (total soil Zn 5.4%, pH 7.3) collected near a Zn smelter. The value of 92 ± 98 mg kg-1 was obtained for the median effective concentration (EC50) values of 0.01 M KNO3 -extractable Zn using the responses of shoot dry biomass, shoot length, and total pigments. This value falls within the range of 95 ± 46 mg kg-1 reported in other studies for the EC50 values of salt-extractable Zn using field-collected soils. The application of ZnCO3 for spiking was able to mimic foliar Zn concentrations of Lolium species observed in field-collected soils. The effective concentrations of soil Zn obtained in the present study are comparable to those obtained in field-collected soils. Future research should determine effective concentrations of metals using soils spiked with metal-containing compounds that mimic a real source of contamination. Environ Toxicol Chem 2020;39:1790-1796. © 2020 SETAC.

Exogenous 6-benzyladenine application affects root morphology by altering hormone status and gene expression of developing lateral roots in Malus hupehensis.

Malus hupehensis is an extensively used apple rootstock in China. In the current study, M. hupehensis seedlings were treated with exogenous 2.2 µm 6-benzyladenine (6-BA) so as to investigate the mechanism by which 6-BA affects lateral root development. The results indicate that 6-BA treatment promotes elongation and thickening of both root and shoot in M. hupehensis, but reduces the number of lateral roots, as well as reducing the auxin level after 6-BA treatment. Moreover, MhAHK4, MhRR1 and MhRR2 were also significantly up-regulated in response to 6-BA treatment. Expression levels of auxin synthesis- and transport-related genes, such as MhYUCCA6, MhYUCCA10, MhPIN1 and MhPIN2, were down-regulated, which corresponds with lower auxin levels in the 6-BA-treated seedlings. A negative regulator of auxin, MhIAA3, was induced by 6-BA treatment, leading to reduced expression of MhARF7 and MhARF19 in 6-BA-treated seedlings. As a result, expression of MhWOX11, MhWOX5, MhLBD16 and MhLBD29 was blocked, which in turn inhibited lateral root initiation. In addition, a lower auxin level decreased expression of MhRR7 and MhRR15, which repressed expression of key transcription factors associated with root development, thus inhibiting lateral root development. In contrast, 6-BA treatment promoted secondary growth (thickening) of the root by inducing expression of MhCYCD3;1 and MhCYCD3;2. Collectively, the changes in hormone levels and gene expression resulted in a reduced number of lateral roots and thicker roots in 6-BA-treated plants.

A genetic framework for regulation and seasonal adaptation of shoot architecture in hybrid aspen.

Shoot architecture is critical for optimizing plant adaptation and productivity. In contrast with annuals, branching in perennials native to temperate and boreal regions must be coordinated with seasonal growth cycles. How branching is coordinated with seasonal growth is poorly understood. We identified key components of the genetic network that controls branching and its regulation by seasonal cues in the model tree hybrid aspen. Our results demonstrate that branching and its control by seasonal cues is mediated by mutually antagonistic action of aspen orthologs of the flowering regulators TERMINAL FLOWER 1 (TFL1) and APETALA1 (LIKE APETALA 1/LAP1). LAP1 promotes branching through local action in axillary buds. LAP1 acts in a cytokinin-dependent manner, stimulating expression of the cell-cycle regulator AIL1 and suppressing BRANCHED1 expression to promote branching. Short photoperiod and low temperature, the major seasonal cues heralding winter, suppress branching by simultaneous activation of TFL1 and repression of the LAP1 pathway. Our results thus reveal the genetic network mediating control of branching and its regulation by environmental cues facilitating integration of branching with seasonal growth control in perennial trees.

Active Vision and Surface Reconstruction for 3D Plant Shoot Modelling.

Plant phenotyping is the quantitative description of a plant's physiological, biochemical, and anatomical status which can be used in trait selection and helps to provide mechanisms to link underlying genetics with yield. Here, an active vision- based pipeline is presented which aims to contribute to reducing the bottleneck associated with phenotyping of architectural traits. The pipeline provides a fully automated response to photometric data acquisition and the recovery of three-dimensional (3D) models of plants without the dependency of botanical expertise, whilst ensuring a non-intrusive and non-destructive approach. Access to complete and accurate 3D models of plants supports computation of a wide variety of structural measurements. An Active Vision Cell (AVC) consisting of a camera-mounted robot arm plus combined software interface and a novel surface reconstruction algorithm is proposed. This pipeline provides a robust, flexible, and accurate method for automating the 3D reconstruction of plants. The reconstruction algorithm can reduce noise and provides a promising and extendable framework for high throughput phenotyping, improving current state-of-the-art methods. Furthermore, the pipeline can be applied to any plant species or form due to the application of an active vision framework combined with the automatic selection of key parameters for surface reconstruction.

Unusual developmental morphology and anatomy of vegetative organs in Utricularia dichotoma-leaf, shoot and root dynamics.

The terrestrial carnivorous species Utricularia dichotoma is known for a great phenotypic plasticity and unusual vegetative organs. Our investigation on 22 sources/populations revealed that after initiation of a leaf and two bladders on a stolon, a bud was formed in the proximal axil of the leaf, developing into a rosette with up to seven organs. The first two primordia of the bud grew into almost every possible combination of organs, but often into two anchor stolons. The patterns were generally not population specific. The interchangeability of organs increased with increasing rank in the succession of organs on stolon nodes. A high potential of switching developmental programs may be successful in a fluctuating environment. In this respect, we were able to show that bladders developed from anchor stolons experimentally when raising the water table. Anatomical structures were simple, lacunate and largely homogenous throughout all organs. They showed similarities with many hydrophytes, reflecting the plant's adaptation to (temporarily) submerged conditions. The principal component analysis was used in the context of dynamic morphology to illustrate correlations between organ types in the morphospace of U. dichotoma, revealing an organ specific patchwork of developmental processes for typical leaves and shoots, and less pronounced for a typical root. The concept and methods we applied may prove beneficial for future studies on the evolution of Lentibulariaceae, and on developmental morphology and genetics of unusual structures in plants.