New approaches for studying and exploiting an old protuberance, the plant trichome

BACKGROUND AND AIMS: Much recent study of plant trichomes has focused on various aspects of glandular secreting trichomes (GSTs) and differentiation of simple trichomes. This Botanical Briefing will highlight: research on various aspects of, and manipulation of glandular secreting trichomes; molecular aspects of the differentiation and development of simple trichomes of arabidopsis and cotton; how methods for manipulation of model systems used in the above work can be applied to expand our understanding of less studied surface structures of plants. SCOPE: The Briefing will cover: established and suggested roles of simple and glandular secreting trichomes; recent results regarding solute and ion movement in trichomes; methods for isolating trichomes; recent studies of trichome differentiation and development; attempts to modify metabolism in secreting trichomes; efforts to exploit trichomes for commercial and agronomic purposes.     

Potassium carrier TRH1 is required for auxin transport in Arabidopsis roots

Disruption of the TRH1 potassium transporter impairs root hair development in Arabidopsis, and also affects root gravitropic behaviour. Rescue of these morphological defects by exogenous auxin indicates a link between TRH1 activity and auxin transport. In agreement with this hypothesis, the rate of auxin translocation from shoots to roots and efflux of [3H]IAA in isolated root segments were reduced in the trh1 mutant, but efflux of radiolabelled auxin was accelerated in yeast cells transformed with the TRH1 gene. In roots, Pro(TRH1):GUS expression was localized to the root cap cells which are known to be the sites of gravity perception and are central for the redistribution of auxin fluxes. Consistent with these findings, auxin-dependent DR5:GUS promoter-reporter construct was misexpressed in the trh1 mutant indicating that partial block of auxin transport through the root cap is associated with upstream accumulation of the phytohormone in protoxylem cells. When [K+] in the medium was reduced from 20 to 0.1 mm, wild type roots showed mild agravitropic phenotype and DR5:GUS misexpression in stelar cells. This pattern of response to low external [K+] was also affected by trh1 mutation. We conclude that the TRH1 carrier is an important part of auxin transport system in Arabidopsis roots.     

The Arabidopsis COW1 gene encodes a phosphatidylinositol transfer protein essential for root hair tip growth

Root hairs are a major site for the uptake of water and nutrients into plants, and they form an increasingly important model system for the study of development in higher plants. We now report on the molecular genetic analysis of the srh1 mutant in Arabidopsis thaliana impaired in root hair tip growth. We show that srh1 is a new allele of cow1 (can of worms1) and we identified the COW1 gene using a positional cloning strategy. The N-terminus of the COW1 protein is 32% identical to an essential phosphatidylinositol transfer protein (PITP), the yeast Sec14 protein (sec14p) while the C-terminus is 34.5% identical to a late nodulin of Lotus japonicus, Nlj16. We show that expression of the COW1 lipid-binding domain complements the growth defect associated with Sec14p dysfunction in yeast. In addition, we show that GFP fused to the COW1 protein specifically accumulates at the site of root hair outgrowth. We conclude that the COW1 protein is a PITP, essential for proper root hair growth.     

Consequences of scavenging behaviour in a plant bug associated with a glandular plant

To elucidate the specific associations of insects with glandular plants, the relationship between a plant bug and a wild azalea was examined. The plant bug Orthotylus gotoi Yasunaga (Hemiptera: Miridae) was found only on shrubs of Rhododendron macrosepalum Maxim. (Ericaceae) in Kyoto, central Japan. R. macrosepalum has leaves, stems, and sepals with dense stalked glands that trap and kill many arthropods. However, nymphs and adults of O. gotoi are able to move freely on R .  macrosepalum . O. gotoi frequently fed on carcasses of various arthropods trapped on R .  macrosepalum , whereas it rarely fed upon plant tissues. In laboratory experiments, O. gotoi nymphs mainly fed on the carcasses of insects, which resulted in enhanced rates of growth and metamorphosis, and adult O. gotoi lived longer when they were provided with insect carcasses. These observations and experiments suggest that O. gotoi is a scavenger. Therefore, if O. gotoi mainly feeds on carcasses of arthropods, this may result in a specific dependence on the glandular plant.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 593–602.     

Glandular trichomes as a barrier against atmospheric oxidative stress: Relationships with ozone uptake,leaf damage,and emission of LOX products across a diverse set of species

There is a spectacular variability in trichome types and densities and trichome metabolites across species, but the functional implications of this variability in protecting from atmospheric oxidative stresses remain poorly understood. The aim of this study was to evaluate the possible protective role of glandular and non‐glandular trichomes against ozone stress. We investigated the interspecific variation in types and density of trichomes and how these traits were associated with elevated ozone impacts on visible leaf damage, net assimilation rate, stomatal conductance, chlorophyll fluorescence, and emissions of lipoxygenase pathway products in 24 species with widely varying trichome characteristics and taxonomy. Both peltate and capitate glandular trichomes played a critical role in reducing leaf ozone uptake, but no impact of non‐glandular trichomes was observed. Across species, the visible ozone damage varied 10.1‐fold, reduction in net assimilation rate 3.3‐fold, and release of lipoxygenase compounds 14.4‐fold, and species with lower glandular trichome density were more sensitive to ozone stress and more vulnerable to ozone damage compared to species with high glandular trichome density. These results demonstrate that leaf surface glandular trichomes constitute a major factor in reducing ozone toxicity and function as a chemical barrier that neutralizes the ozone before it enters the leaf.     

Monte Carlo evaluation of glandular dose in cone-beam X-ray computed tomography dedicated to the breast: Homogeneous and heterogeneous breast models

PurposeIn cone-beam computed tomography dedicated to the breast (BCT), the mean glandular dose (MGD) is the dose metric of reference, evaluated from the measured air kerma by means of normalized glandular dose coefficients (DgN_CT). This work aimed at computing, for a simple breast model, a set of DgN_CT values for monoenergetic and polyenergetic X-ray beams, and at validating the results vs. those for patient specific digital phantoms from BCT scans.MethodsWe developed a Monte Carlo code for calculation of monoenergetic DgN_CT coefficients (energy range 4.25–82.25 keV). The pendant breast was modelled as a cylinder of a homogeneous mixture of adipose and glandular tissue with glandular fractions by mass of 0.1%, 14.3%, 25%, 50% or 100%, enveloped by a 1.45 mm-thick skin layer. The breast diameter ranged between 8 cm and 18 cm. Then, polyenergetic DgN_CT coefficients were analytically derived for 49-kVp W-anode spectra (half value layer 1.25–1.50 mm Al), as in a commercial BCT scanner. We compared the homogeneous models to 20 digital phantoms produced from classified 3D breast images.ResultsPolyenergetic DgN_CT resulted 13% lower than most recent published data. The comparison vs. patient specific breast phantoms showed that the homogeneous cylindrical model leads to a DgN_CT percentage difference between −15% and +27%, with an average overestimation of 8%.ConclusionsA dataset of monoenergetic and polyenergetic DgN_CT coefficients for BCT was provided. Patient specific breast models showed a different volume distribution of glandular dose and determined a DgN_CT 8% lower, on average, than homogeneous breast model.     

Supravital Methylene Blue Staining of Piloneural Complexes of Common Fur Hair Follicles in the Rat

Light microscopic observations employing supravital methylene blue staining are presented for piloneural complexes of common fur hairs in the mystacial pad of the rat snout. The investigation revealed anatomical details of piloneural complexes belonging to follicles of both vellus and guard hairs. In the methylene blue stained preparations, different types of palisade-like lanceolate nerve fiber endings could be discriminated. The thicker vellus and thinner guard hairs (hair diameter: 15-25 μm) exhibited a different innervation pattern compared to the thicker guard hairs, and two subtypes of piloneural complexes could be distinguished. Both subtypes were characterized by slightly stained lanceolate endings and the absence of a circular nerve fiber plexus. One subtype, however, showed strongly stained spines originating from the lanceolate endings. A few spines of adjacent lanceolate endings appeared in contact with each other. In the second subtype, these spines were replaced by anastomoses suggesting a delicate terminal nerve fiber network. The moderately stained lanceolate endings located primarily at the follicles of thicker guard hairs (hair diameter: 30-40 μm) showed smooth outlines, but were characterized by the occurrence of an intensely stained additional circular nerve fiber plexus. The differences in the morphology of piloneural complexes associated with the follicles of common fur hairs suggest differences regarding their mechanoreceptive tasks.     

Developmental plasticity of glandular trichomes into somatic embryogenesis in Tilia amurensis

BACKGROUND AND AIMS: In Tilia amurensis, two types of trichomes (hairy and glandular) develop from epidermal surfaces of cotyledons and hypocotyls of zygotic embryos soon after germination. Here, it is demonstrated that glandular trichome initials develop directly into somatic embryos when treated in vitro with 2,4-dichlorophenoxyacetic acid (2,4-D). METHODS: Zygotic embryos of Tilia amurensis were cultured on Murashige and Skoog medium with 3 % sucrose and various concentrations (0, 2.2, 4.4 and 8.8 microm) of 2,4-D. Morphological development of trichomes and somatic embryos was analysed by scanning electron microscope and light microscope after histological sectioning. KEY RESULTS: In zygotic embryos cultured on medium with 4.4 microM 2,4-D, formation of hairy trichomes was completely suppressed but formation of glandular trichome initials increased. That some filamentous trichome initials developed directly into somatic embryos was confirmed by histological and scanning electron microscope observation. When explants with different stages of trichome initials (two-, four- and eight-celled filamentous and fully mature trichomes) were temporally pre-treated with 4.4 microM 2,4-D for 24 h and transferred into hormone-free medium, two-celled and four-celled filamentous trichome initials were the effective stage of trichomes for somatic embryo induction. CONCLUSIONS: It is suggested that early developing filamentous trichome initials have developmental plasticity and that with 2,4-D treatment these trichome initials develop directly into somatic embryos.     

Phosphorus (P) uptake and growth of a root hairless barley mutant (bald root barley, brb) and wild type in low- and high-P soils

The recently isolated root‐hairless mutant of barley (Hordeum vulgare L), bald root barley, brb offers a unique possibility to quantify the importance of root hairs in phosphorus (P) uptake from soil. In the present study the ability of brb and the wild‐type, barley genotype Pallas producing normal root hairs to deplete P in the rhizosphere soil was investigated and the theory of diffusion and mass flow applied to compare the predicted and measured depletion profiles of diffusible P. Pallas depleted twice as much P from the rhizosphere soil as brb. The P depletion profile of Pallas uniformly extended to 0.8 mm from the root surface, which was equal to the root hair length (RHL). The model based on the theory of diffusion and mass flow explained the observed P‐depletion profile of brb, and the P depletion outside the root‐hair zone of Pallas, suggesting that the model is valid only for P movement in rhizosphere soil outside the root‐hair zone. In low‐P soil (P in soil solution 3 µm ) brb did not survive after 30 d, whereas Pallas continued to grow, confirming the importance of root hairs in plant growth in a P‐limiting environment. In high‐P soil (P in soil solution 10 µm ) both brb and Pallas maintained their growth, and they were able to produce seeds. At the high‐P concentration, RHL of the Pallas was reduced from 0.80 ± 0.2 to 0.68 ± 0.14 mm. In low‐P soil, P‐uptake rate into the roots of Pallas was 4.0 × 10^?7 g mm^?1 d^?1 and that of brb was 1.9 × 10^?7 g mm^?1 d^?1, which agreed well with the double amount of P depleted from the rhizosphere soil of Pallas in comparison with that of brb. In high‐P soil, the P uptake rates into the roots of brb and Pallas were 3.3 and 5.5 × 10^?7 g mm^?1 d^?1, respectively. The results unequivocally confirmed that in a low‐P environment, root hairs are of immense importance in P acquisition and plants survival, but under high‐P conditions they may be dispensable. The characterization of phenotypes brb and Pallas and the ability to reproduce seeds offers a unique possibility of molecular mapping of QTLs and candidate genes conferring root‐hair formation and growth of barley.