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Arabidopsis (Arabidopsis thaliana) leaf trichomes are single-cell structures with a well-studied development, but little is understood about their function. Developmental studies focused mainly on the early shaping stages, and little attention has been paid to the maturation stage. We focused on the EXO70H4 exocyst subunit, one of the most up-regulated genes in the mature trichome. We uncovered EXO70H4-dependent development of the secondary cell wall layer, highly autofluorescent and callose rich, deposited only in the upper part of the trichome. The boundary is formed between the apical and the basal parts of mature trichome by a callose ring that is also deposited in an EXO70H4-dependent manner. We call this structure the Ortmannian ring (OR). Both the secondary cell wall layer and the OR are absent in the exo70H4 mutants. Ecophysiological aspects of the trichome cell wall thickening include interference with antiherbivore defense and heavy metal accumulation. Ultraviolet B light induces EXO70H4 transcription in a CONSTITUTIVE PHOTOMORPHOGENIC1-dependent way, resulting in stimulation of trichome cell wall thickening and the OR biogenesis. EXO70H4-dependent trichome cell wall hardening is a unique phenomenon, which may be conserved among a variety of the land plants. Our analyses support a concept that Arabidopsis trichome is an excellent model to study molecular mechanisms of secondary cell wall deposition.
- MeSH
- Arabidopsis metabolismus účinky záření ultrastruktura MeSH
- buněčná stěna metabolismus ultrastruktura MeSH
- cytokineze účinky záření MeSH
- fluorescence MeSH
- glukany metabolismus MeSH
- měď metabolismus MeSH
- mutace genetika MeSH
- podjednotky proteinů metabolismus MeSH
- proteiny huseníčku metabolismus MeSH
- trichomy metabolismus účinky záření ultrastruktura MeSH
- ultrafialové záření MeSH
- vezikulární transportní proteiny metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Názvy látek
- callose MeSH Prohlížeč
- EXO70H4 protein, Arabidopsis MeSH Prohlížeč
- glukany MeSH
- měď MeSH
- podjednotky proteinů MeSH
- proteiny huseníčku MeSH
- vezikulární transportní proteiny MeSH
BACKGROUND AND AIMS: Root hemiparasites from the rhinanthoid clade of Orobanchaceae possess metabolically active glandular trichomes that have been suggested to function as hydathode trichomes actively secreting water, a process that may facilitate resource acquisition from the host plant's root xylem. However, no direct evidence relating the trichomes to water secretion exists, and carbon budgets associated with this energy-demanding process have not been determined. METHODS: Macro- and microscopic observations of the leaves of hemiparasitic Rhinanthus alectorolophus were conducted and night-time gas exchange was measured. Correlations were examined among the intensity of guttation, respiration and transpiration, and analysis of these correlations allowed the carbon budget of the trichome activity to be quantified. We examined the intensity of guttation, respiration and transpiration, correlations among which indicate active water secretion. KEY RESULTS: Guttation was observed on the leaves of 50 % of the young, non-flowering plants that were examined, and microscopic observations revealed water secretion from the glandular trichomes present on the abaxial leaf side. Night-time rates of respiration and transpiration and the presence of guttation drops were positively correlated, which is a clear indicator of hydathode trichome activity. Subsequent physiological measurements on older, flowering plants indicated neither intense guttation nor the presence of correlations, which suggests that the peak activity of hydathodes is in the juvenile stage. CONCLUSIONS: This study provides the first unequivocal evidence for the physiological role of the hydathode trichomes in active water secretion in the rhinanthoid Orobanchaceae. Depending on the concentration of organic elements calculated to be in the host xylem sap, the direct effect of water secretion on carbon balance ranges from close to neutral to positive. However, it is likely to be positive in the xylem-only feeding holoparasites of the genus Lathraea, which is closely related to Rhinanthus. Thus, water secretion by the hydathodes might be viewed as a physiological pre-adaptation in the evolution of holoparasitism in the rhinanthoid lineage of Orobanchaceae.
- Klíčová slova
- Ecophysiology, Lathraea, Rhinanthus alectorolophus, Triticum aestivum, holoparasite, hydathode trichome, orobanche, parasitic plant, respiration, rhinanthoid Orobanchaceae, root hemiparasite, transpiration, water regime, water secretion, xylem,
- MeSH
- biologická evoluce * MeSH
- buněčné dýchání MeSH
- interakce hostitele a parazita fyziologie MeSH
- kořeny rostlin parazitologie MeSH
- lineární modely MeSH
- listy rostlin metabolismus MeSH
- Orobanchaceae fyziologie MeSH
- plyny metabolismus MeSH
- transpirace rostlin MeSH
- trichomy fyziologie MeSH
- voda metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Názvy látek
- plyny MeSH
- voda MeSH
Almost all dried vegetative trichomes ofAnabaena iyengarii, Westiellopsis prolifica andNostochopsis lobatus died within 1 h, while those ofOscillatoria acuminata retained viability to some extent for 1 d under similar storage conditions. The viability of dried vegetative trichomes ofO. acuminata decreased about equally on storage at 20 degrees C in the light or in the dark, but dropped rapidly at 12 and 0 degrees C in the dark. Vegetative trichomes ofA. iyengarii, N. lobatus andW. prolifica were more sensitive to frost than those ofO. acuminata, and this correlated with their low resistance to desiccation because both types of exposure involved osmotic stress. Both dried and wet akinetes ofA. iyengarii, W. prolifica andN. lobatus were about equally viable when stored at 20 degrees C in the light or the dark or at 12 and 0 degrees C in the dark, but their germination ability decreased on storage at 0 degrees C. The water stress imposed on growing vegetative trichomes either in high-agar media or in NaCl-supplemented liquid media reduced the survival ofO. acuminata trichomes, decreased or totally suppressed akinete and heterocyst formation and akinete germination inA. iyengarii, W. prolifica andN. lobatus. The sensitivity decreased in the sequenceA. iyengarii
- Publikační typ
- časopisecké články MeSH
The Rhinanthoid clade of the family Orobanchaceae comprises plants displaying a hemiparasitic or holoparasitic strategy of resource acquisition. Some of its species (mainly Rhinanthus spp.) are often used as models for studies of hemiparasite physiology. Although there is a well-developed concept covering their physiological processes, most recent studies have neglected the existence of hydathode trichomes present on leaves of these hemiparasitic plants. As a first step for the proposed integration of these structures in the theory of physiological processes of the hemiparasites, we described the outer micromorphology and ultrastructure of the hydathode trichomes on leaves of hemiparasitic Rhinanthus alectorolophus and Odontites vernus with scanning and transmission electron microscopy (SEM and TEM, respectively). The TEM inspections of both types of trichome revealed typical ultrastructural features: labyrinthine cell wall, high content of cytoplasm in cells with numerous mitochondria and presence of plasmodesmata. All these features indicate high metabolic activity complying with their function as glandular trichomes actively secreting water. The active secretion of water by the hydathode trichomes (evidence for which is summarised here) also presents a possible mechanism explaining results of previous gas exchange measurements detecting high dark respiration and transpiration rates and a tight inter-correlation between them in hemiparasitic Orobanchaceae. In addition, this process is hypothesised to have allowed multiple evolutionary transitions from facultative to obligate hemiparasitism and unique xylem-feeding holoparasitism of Lathraea with a long-lived underground stage featuring a rhizome covered by scales of leaf origin.
- MeSH
- biologická evoluce MeSH
- buněčná stěna ultrastruktura MeSH
- buněčné dýchání MeSH
- epidermis rostlin genetika fyziologie ultrastruktura MeSH
- listy rostlin genetika fyziologie ultrastruktura MeSH
- mikroskopie elektronová rastrovací MeSH
- mitochondrie ultrastruktura MeSH
- Orobanchaceae genetika fyziologie ultrastruktura MeSH
- plazmodesmy ultrastruktura MeSH
- transmisní elektronová mikroskopie MeSH
- transpirace rostlin MeSH
- voda metabolismus MeSH
- xylém genetika fyziologie ultrastruktura MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Názvy látek
- voda MeSH
Exocyst component of 70-kDa (EXO70) proteins are constituents of the exocyst complex implicated in vesicle tethering during exocytosis. MILDEW RESISTANCE LOCUS O (MLO) proteins are plant-specific calcium channels and some MLO isoforms enable fungal powdery mildew pathogenesis. We here detected an unexpected phenotypic overlap of Arabidopsis thaliana exo70H4 and mlo2 mlo6 mlo12 triple mutant plants regarding the biogenesis of leaf trichome secondary cell walls. Biochemical and Fourier transform infrared spectroscopic analyses corroborated deficiencies in the composition of trichome cell walls in these mutants. Transgenic lines expressing fluorophore-tagged EXO70H4 and MLO exhibited extensive colocalization of these proteins. Furthermore, mCherry-EXO70H4 mislocalized in trichomes of the mlo triple mutant and, vice versa, MLO6-GFP mislocalized in trichomes of the exo70H4 mutant. Expression of GFP-marked PMR4 callose synthase, a known cargo of EXO70H4-dependent exocytosis, revealed reduced cell wall delivery of GFP-PMR4 in trichomes of mlo triple mutant plants. In vivo protein-protein interaction assays in plant and yeast cells uncovered isoform-preferential interactions between EXO70.2 subfamily members and MLO proteins. Finally, exo70H4 and mlo6 mutants, when combined, showed synergistically enhanced resistance to powdery mildew attack. Taken together, our data point to an isoform-specific interplay of EXO70 and MLO proteins in the modulation of trichome cell wall biogenesis and powdery mildew susceptibility.
- MeSH
- Arabidopsis * metabolismus MeSH
- buněčná stěna metabolismus MeSH
- nemoci rostlin mikrobiologie MeSH
- odolnost vůči nemocem genetika MeSH
- protein - isoformy genetika metabolismus MeSH
- proteiny huseníčku * genetika metabolismus MeSH
- rostlinné proteiny metabolismus MeSH
- trichomy genetika metabolismus MeSH
- vezikulární transportní proteiny metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- Názvy látek
- EXO70H4 protein, Arabidopsis MeSH Prohlížeč
- protein - isoformy MeSH
- proteiny huseníčku * MeSH
- rostlinné proteiny MeSH
- vezikulární transportní proteiny MeSH
In Utricularia, the flower spur is a nectary and in this organ, nectar is produced and stored. This study aimed to examine the structure of the nectary trichomes in four Utricularia species (Utricularia vulgaris L., U. australis R.Br., U. bremii Heer and U. foliosa L.) from the generic section Utricularia. We have investigated whether species with different spur morphology had similar spur anatomy and nectary trichome structure. In Utricularia flowers, nectar is produced by spur capitate trichomes (sessile or stalked). Our results showed that regardless of the various spur morphology, trichomes have similar architecture and ultrastructure. Head cells of these trichomes are transfer cells with an eccrine nectar secretion. Examined species differed in the micromorphology of papillae in spurs. The fly Eristalis tenax was found to be a pollinator of U. vulgaris. Small Halictidae bees seem to be pollinators of U. foliosa.
- Klíčová slova
- Bladderwort, Carnivorous plant, Entomophily, Floral micromorphology, Lentibulariaceae, Nectar, Nectary ultrastructure, Pollination, Sect. Utricularia, Spur, Trichomes,
- MeSH
- květy chemie MeSH
- opylení imunologie MeSH
- rostlinný nektar chemie MeSH
- Publikační typ
- časopisecké články MeSH
- Názvy látek
- rostlinný nektar MeSH
Plasma membrane (PM) lipid composition and domain organization are modulated by polarized exocytosis. Conversely, targeting of secretory vesicles at specific domains in the PM is carried out by exocyst complexes, which contain EXO70 subunits that play a significant role in the final recognition of the target membrane. As we have shown previously, a mature Arabidopsis trichome contains a basal domain with a thin cell wall and an apical domain with a thick secondary cell wall, which is developed in an EXO70H4-dependent manner. These domains are separated by a cell wall structure named the Ortmannian ring. Using phospholipid markers, we demonstrate that there are two distinct PM domains corresponding to these cell wall domains. The apical domain is enriched in phosphatidic acid (PA) and phosphatidylserine, with an undetectable amount of phosphatidylinositol 4,5-bisphosphate (PIP2), whereas the basal domain is PIP2-rich. While the apical domain recruits EXO70H4, the basal domain recruits EXO70A1, which corresponds to the lipid-binding capacities of these two paralogs. Loss of EXO70H4 results in a loss of the Ortmannian ring border and decreased apical PA accumulation, which causes the PA and PIP2 domains to merge together. Using transmission electron microscopy, we describe these accumulations as a unique anatomical feature of the apical cell wall-radially distributed rod-shaped membranous pockets, where both EXO70H4 and lipid markers are immobilized.
- Klíčová slova
- EXO70, cell wall, exocyst complex, phosphatidic acid, phosphatidylinositol 4,5-bisphosphate, phospholipids, plasma membrane domains, polar exocytosis, trichome,
- MeSH
- Arabidopsis chemie genetika MeSH
- buněčná membrána chemie genetika MeSH
- exocytóza genetika MeSH
- fosfatidylinositol-4,5-difosfát chemie metabolismus MeSH
- fosfatidylseriny chemie genetika MeSH
- membránové lipidy genetika metabolismus MeSH
- proteiny huseníčku chemie genetika MeSH
- trichomy chemie genetika MeSH
- vezikulární transportní proteiny chemie genetika MeSH
- Publikační typ
- časopisecké články MeSH
- Názvy látek
- EXO70A1 protein, Arabidopsis MeSH Prohlížeč
- EXO70H4 protein, Arabidopsis MeSH Prohlížeč
- fosfatidylinositol-4,5-difosfát MeSH
- fosfatidylseriny MeSH
- membránové lipidy MeSH
- proteiny huseníčku MeSH
- vezikulární transportní proteiny MeSH
There is an enormous diversity in the structure of the flower palate of the carnivorous rootless genus Utricularia. This study aims to examine the structure of the palates in Utricularia bremii Heer and U. minor L of the Utricularia sect. Utricularia, which have a glandular palate type. In both species, the palate has only one type of glandular trichomes. Because of the occurrence of cell wall ingrowths in its glandular cells, any exudation may be transported via eccrinous secretion. It was proposed that the palate trichomes of the examined species act as scent glands and that the palate may play a role as an unguentarium. Both U. bremii and U. minor are of an open flower type. Thus, U. bremii and U. minor flowers can be penetrated by small, weak insects, which then easily have access to their generative structure. Small Hymenoptera (member of families Mymaridae and Braconidae) were observed as flower visitors of the male-sterile species Utricularia bremii.
- Klíčová slova
- Bladderwort, Carnivorous plant, Floral micro-morphology, Lentibulariaceae, Osmophore, Pollination, Sect. Utricularia, Ultrastructure,
- MeSH
- fylogeneze MeSH
- hmyz fyziologie MeSH
- květy metabolismus fyziologie MeSH
- Magnoliopsida anatomie a histologie metabolismus MeSH
- trichomy anatomie a histologie metabolismus MeSH
- zvířata MeSH
- Check Tag
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
Viroids are small infectious pathogens, composed of a short single-stranded circular RNA. Hop (Humulus lupulus L.) plants are hosts to four viroids from the family Pospiviroidae. Hop latent viroid (HLVd) is spread worldwide in all hop-growing regions without any visible symptoms on infected hop plants. In this study, we evaluated the influence of HLVd infection on the content and the composition of secondary metabolites in maturated hop cones, together with gene expression analyses of involved biosynthesis and regulation genes for Saaz, Sládek, Premiant and Agnus cultivars. We confirmed that the contents of alpha bitter acids were significantly reduced in the range from 8.8% to 34% by viroid infection. New, we found that viroid infection significantly reduced the contents of xanthohumol in the range from 3.9% to 23.5%. In essential oils of Saaz cultivar, the contents of monoterpenes, terpene epoxides and terpene alcohols were increased, but the contents of sesquiterpenes and terpene ketones were decreased. Secondary metabolites changes were supported by gene expression analyses, except essential oils. Last-step biosynthesis enzyme genes, namely humulone synthase 1 (HS1) and 2 (HS2) for alpha bitter acids and O-methytransferase 1 (OMT1) for xanthohumol, were down-regulated by viroid infection. We found that the expression of ribosomal protein L5 (RPL5) RPL5 and the splicing of transcription factor IIIA-7ZF were affected by viroid infection and a disbalance in proteosynthesis can influence transcriptions of biosynthesis and regulatory genes involved in of secondary metabolites biosynthesis. We suppose that RPL5/TFIIIA-7ZF regulatory cascade can be involved in HLVd replication as for other viroids of the family Pospiviroidae.
- Klíčová slova
- HLVd, Humulus lupulus, bitter acids content, differential gene expression, essential oils, hop, hop latent viroid, xanthohumol,
- Publikační typ
- časopisecké články MeSH
Escalation (macroevolutionary increase) or divergence (disparity between relatives) in trait values are two frequent outcomes of the plant-herbivore arms race. We studied the defences and caterpillars associated with 21 sympatric New Guinean figs. Herbivore generalists were concentrated on hosts with low protease and oxidative activity. The distribution of specialists correlated with phylogeny, protease and trichomes. Additionally, highly specialised Asota moths used alkaloid rich plants. The evolution of proteases was conserved, alkaloid diversity has escalated across the studied species, oxidative activity has escalated within one clade, and trichomes have diverged across the phylogeny. Herbivore specificity correlated with their response to host defences: escalating traits largely affected generalists and divergent traits specialists; but the effect of escalating traits on extreme specialists was positive. In turn, the evolution of defences in Ficus can be driven towards both escalation and divergence in individual traits, in combination providing protection against a broad spectrum of herbivores.
- Klíčová slova
- Alkaloids, Choreutidae, Lepidoptera, New Guinea, Pyraloidea, coevolution, cysteine protease, herbivore, polyphenols, trichomes,
- MeSH
- býložravci * MeSH
- fenotyp MeSH
- Ficus * MeSH
- fylogeneze MeSH
- hmyz * MeSH
- zvířata MeSH
- Check Tag
- zvířata MeSH
- Publikační typ
- dopisy MeSH