Growth and ultrastructure ofArabidopsis root hairs: therhd3 mutation alters vacuole enlargement and tip growth

The root hairs of plants are tubular projections of root epidermal cells and are suitable for investigating the control of cellular morphogenesis. In wild-typeArabidopsis thaliana (L.) Heynh, growing root hairs were found to exhibit cellular expansion limited to the apical end of the cell, a polarized distribution of organelles in the cytoplasm, and vesicles of several types located near the growing tip. Therhd3 mutant produces short and wavy root hairs with an average volume less than one-third of the wild-type hairs, indicating abnormal cell expansion. The mutant hairs display a striking reduction in vacuole size and a corresponding increase in the relative proportion of cytoplasm throughout hair development. Bead-labeling experiments and ultrastructural analyses indicate that the wavy-hair phenotype of the mutant is caused by asymmetric tip growth, possibly due to abnormally distributed vesicles in cortical areas flanking the hair tips. It is suggested that a major effect of therhd3 mutation is to inhibit vacuole enlargement which normally accompanies root hair cell expansion.     

Variable responses of tuber moth to the leaf trichomes of wild potatoes

This study examines the response of tuber moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), during the initial stages of attack, to variability in trichome density and composition on foliage of Solanum berthaultii (Hawkes) and Solanum tarijense (Hawkes) (Solanaceae). Solanum berthaultii bears two types of glandular trichome (type A and type B) that together reduced oviposition by the moth. Females were often completely deterred from ovipositing on foliage with >300 trichomes per cm². In contrast, neonate establishment on S. berthaultii was generally positively related to trichome densities, indicating that trichomes may be a poor defense against P. operculella when the moth oviposits in soil and neonate larvae select the host plant. Solanum tarijense has only one type of glandular trichome (type A) and eglandular hairs. Most eggs were deposited on the adaxial leaf surfaces that had lower trichome densities. Although the density of type A trichomes was negatively related to oviposition, high densities of hairs on the abaxial and adaxial leaf surfaces appeared to stimulate oviposition, leading to stronger positive relations between hair densities and oviposition. Larvae generally established on the abaxial surface where hair densities were greatest. Relationships between the abaxial densities of leaf hairs and neonate establishment on S. tarijense were positive. The results indicate that the responses by P. operculella to the types and density of trichomes are complex. Whereas type A and type B trichomes may act synergistically to reduce oviposition by the moth, leaf hairs do not defend against oviposition and neither leaf hairs nor type A and B trichomes reduce neonate establishment by this herbivore species.     

Etude infrastructurale de la stipule de Vicia faba L. au niveau du nectaire

Summary In the extrafloral nectary of the broad bean there is evidence of two fundamental types of cells: one with dense hyaloplasm, well developed ergastoplasm and golgi apparatus, all features of glandular cells, and another with opposite features. The cells of the head of the secretory hairs and those of the subjacent epidermis which are not prolonged with such a hair are of the first type. The epidermal cells prolonged with a hair and the pedicellar cell of this hair are of the second type. Moreover, the companion cells of the subjacent conducting bundle look like cells of the first type, especially those of the head of the secretory hairs owing to their numerous wall protuberances. Cells of the second type are presumably involved in transit processes between phloem and trichome, and cells of the first type in excretory processes.

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Ce travail fait partie d'une Thèse de Doctorat d'Etat sur la cyto-physiologie des nectaires. — (Travail effectué au Laboratoire de Bot. Appl. et Microbiologie et au Centre de Microscopie électronique de la Faculté des Sciences de Bordeaux (France).     

Nitrate induction of root hair density is mediated by TGA1/TGA4 and CPC transcription factors in Arabidopsis thaliana

Root hairs are specialized cells that are important for nutrient uptake. It is well established that nutrients such as phosphate have a great influence on root hair development in many plant species. Here we investigated the role of nitrate on root hair development at a physiological and molecular level. We showed that nitrate increases root hair density in Arabidopsis thaliana. We found that two different root hair defective mutants have significantly less nitrate than wild‐type plants, suggesting that in A. thaliana root hairs have an important role in the capacity to acquire nitrate. Nitrate reductase‐null mutants exhibited nitrate‐dependent root hair phenotypes comparable with wild‐type plants, indicating that nitrate is the signal that leads to increased formation of root hairs. We examined the role of two key regulators of root hair cell fate, CPC and WER, in response to nitrate treatments. Phenotypic analyses of these mutants showed that CPC is essential for nitrate‐induced responses of root hair development. Moreover, we showed that NRT1.1 and TGA1/TGA4 are required for pathways that induce root hair development by suppression of longitudinal elongation of trichoblast cells in response to nitrate treatments. Our results prompted a model where nitrate signaling via TGA1/TGA4 directly regulates the CPC root hair cell fate specification gene to increase formation of root hairs in A. thaliana.     

Ultrastructural ontogeny of leaf cavity trichomes inAzolla implies a functional role in metabolite exchange

Summary Anabaena azollae is associated with two types of multicellular epidermal trichomes inAzolla leaf cavities, the simple and branched hairs. The observation of transfer cell ultrastructure in some hair cells led to speculation that the cavity hairs might participate in metabolite exchange between the symbionts. The developmental ontogeny of cavity trichomes is described here, using transmission electron microscopy, with a goal of improving our understanding of possible functions of these structures in the symbiosis. The observations have established that all cells of simple and branched hairs develop the structural characteristics of transfer cells, but not simultaneously. Rather, there is an acropetal succession of transfer cell ultrastructure beginning in terminal cells, moving to body cells where present, and ending in stalk cells. The transfer cell stage is followed immediately by senescence in all hair cells. The timing of transfer cell differentiation, considered together with information from other studies, suggests that branched hairs may be involved in exchange of fixed nitrogen between the symbionts, while simple hairs may participate in exchange of fixed carbon fromAzolla toAnabaena. Contribution no. 869 from the Battelle-C. F. Kettering Research Laboratory.     

3种龙葵表皮毛类型及发育过程观察研究

通过观察发现龙葵（Ｓｏｌａｎｕｍ ｎｉｇｒｕｍ Ｌ．）、少花龙葵（Ｓ．ｐｈｏｔｅｉｎｏｃａｒｐｕｍ Ｎａｋａｍ,ｅｔＯ－ｄｓｈｉ）和黄果龙葵（Ｓ．ｎｉｇｕｍ Ｌ．ｖａｒ．ｓｕａｖｅｏｌｅｎｓ Ｇ．Ｌ．Ｇｕｏ）的表皮毛均为腺毛,主要有单细胞头腺毛和多细胞头腺毛２种。腺毛的原始细胞都来源于原表皮细胞,经２次平周分裂产生基细胞、柄细胞和顶端细胞、在腺毛后期的形态发生中,柄细胞和顶细胞的分裂状态决定腺毛的     

Hair morphology and systematics ofPhysalis (Solanaceae)

Hair morphology was exammed on developing seedlings and mature plants of ten species from five of seven series inPhysalis. The taxonomic importance of hair development and the distribution of hair types then was evaluated with respect to other comparative data. Two classes of hair types were observed, as found in earlier investigations ofSolanum andRhododendron. The first hair class comprises a sequence from unbranched hairs to dendroid-stelliform hairs. Hairs of this class show considerable variability among taxa in size, presence or absence of glandular tips, and presence and degree of branching, and so were useful in making taxonomic comparisons. The second hair class consists of short hairs, each tipped with a multicellular gland. This second hair class was not useful taxonomically because of its lack of variability in morphology and its often rare occurrence on a plant. The taxa studied exhibit distinctive trichome features which serve to distinguish series and, in some cases together with other data, provided new insight into previously enigmatic species relationships.     

X-ray Microanalysis of Ion Distribution in Abutilon Nectary Hairs

Robards, A. W. and Oates, K. 1986. X-ray microanalysis of iondistribution in Abutilon nectaryhairs—J. exp. Bot. 37:940–946. X-ray microanalysis of frozen-hydrated Abutilon nectary hairsshows that potassium is present at easily detectable levelsin all cells between the phloem and the trichome tip cells.There is little overall variation in the amount of potassiumfrom cell to cell although it was found that: (i) some basalcells accumulate potassium to a significantly higher level thanthe hair cells; and (ii) there is more potassium in the cellstowards the distal end of the trichome. These findings wouldbe consistent with a model for secretion envisaging a trans-membraneloading of nectar into a ‘secretory reticulum’ inall trichome cells. This loading process would be selectiveand would exclude ions. A pronounced increase in the level ofchlorine from the basal cell along the hair was observed. Nocomplementary cation was detected but equivalent changes inthe level of Na would have been below the detection limit ofthe system. Key words: Abutilon, nectary hairs, X-ray microanalysis, ion distribution     

Trichome studies in Parthenium hysterophorus and their taxonomic importance

In Parthenium hysterophorus L. four trichome types i.e. (1) cylindrical hair, (2) moniliform hair, (3) simple multiseriate hair and (4) shaggy hair, have been recognized mostly on the basis of their structural differences. In the past, these types have not been observed in any species of Parthenium studied so far. Out of these four, two types i.e. cylindrical and moniliform hairs are known to occur in the Compositae and remaining two i.e. simple multiseriate and shaggy hairs are the new forms for the family and have been observed for the first time in presently investigated species. Trichomes are an important taxonomic tool in distinguishing between genera and species. Trichomes of the different species of Parthenium as noticed from their structure are useful in distinguishing these species and even their corresponding organs.     

Hair density in the Eurasian otter <Emphasis Type="Italic">Lutra lutra</Emphasis> and the Sea otter <Emphasis Type="Italic">Enhydra lutris</Emphasis>

The hair density of adult Eurasian otters Lutra lutra (Linnaeus, 1758) and sea otters Enhydra lutris (Linnaeus, 1758) was analysed using skin samples taken from frozen carcasses. Lutra lutra exhibited a mean hair density of about 70 000 hairs/cm² (whole body, appendages excepted), the mean individual density ranging from about 60 000 to 80 000 hairs/cm². The dominant hair type were secondary hairs (wool hairs), the hair coat comprising only 1.26% of primary hairs (PH). Secondary hair (SH) density remained constant over the body (appendages excepted), whereas a few variations in PH density were observed. Neither an influence of the sex, nor a seasonal variation of the hair coat was found, moulting seems to be continuous. Enhydra lutris had a hair density between 120 000 and 140 000 hairs/cm², the primary hairs representing less than 1% within the hair coat. Hair density remained quite constant over the regions of the trunk but was lower at the head (about 60 000 hairs/cm² on the cheek). The hair follicles were arranged in specific groups with different bundles of varying size, normally comprising dominant numbers of wool hair (SH) follicles. Invariably there was always a large central primary hair follicle and numerous sebaceous glands between the bundles and principally around the PH follicles. The results are discussed related to possible ecological influences on hair coat density.     

Root hair-specific expansins modulate root hair elongation in rice

Root hair growth requires intensive cell‐wall modification. This study demonstrates that root hair‐specific expansin As, a sub‐clade of the cell wall‐loosening expansin proteins, are required for root hair elongation in rice (Oryza sativa L.). We identified a gene encoding EXPA17 (OsEXPA17) from a rice mutant with short root hairs. Promoter::reporter transgenic lines exhibited exclusive OsEXPA17 expression in root hair cells. The OsEXPA17 mutant protein (OsexpA17) contained a point mutation, causing a change in the amino acid sequence (Gly104→Arg). This amino acid alteration is predicted to disrupt a highly conserved disulfide bond in the mutant. Suppression of OsEXPA17 by RNA interference further confirmed requirement for the gene in root hair elongation. Complementation of the OsEXPA17 mutant with other root hair EXPAs (OsEXPA30 and Arabidopsis EXPA7) can restore root hair elongation, indicating functional conservation of these root hair EXPAs in monocots and dicots. These results demonstrate that members of the root hair EXPA sub‐clade play a crucial role in root hair cell elongation in Graminaceae.     

The Ca2+‐binding protein PCaP2 located on the plasma membrane is involved in root hair development as a possible signal transducer

Plasma membrane‐associated Ca²⁺‐binding protein–2 (PCaP2) of Arabidopsis thaliana is a novel‐type protein that binds to the Ca²⁺/calmodulin complex and phosphatidylinositol phosphates (PtdInsPs) as well as free Ca²⁺. Although the PCaP2 gene is predominantly expressed in root hair cells, it remains unknown how PCaP2 functions in root hair cells via binding to ligands. From biochemical analyses using purified PCaP2 and its variants, we found that the N–terminal basic domain with 23 amino acids (N23) is necessary and sufficient for binding to PtdInsPs and the Ca²⁺/calmodulin complex, and that the residual domain of PCaP2 binds to free Ca²⁺. In mutant analysis, a pcap2 knockdown line displayed longer root hairs than the wild‐type. To examine the function of each domain in root hair cells, we over‐expressed PCaP2 and its variants using the root hair cell‐specific EXPANSIN A7 promoter. Transgenic lines over‐expressing PCaP2, PCaP2^G2A (second glycine substituted by alanine) and ?23PCaP2 (lacking the N23 domain) exhibited abnormal branched and bulbous root hair cells, while over‐expression of the N23 domain suppressed root hair emergence and elongation. The N23 domain was necessary and sufficient for the plasma membrane localization of GFP‐tagged PCaP2. These results suggest that the N23 domain of PCaP2 negatively regulates root hair tip growth via processing Ca²⁺ and PtdInsP signals on the plasma membrane, while the residual domain is involved in the polarization of cell expansion.     

Pedestal hairs of the ant Echinopla melanarctos (Hymenoptera, Formicidae): morphology and functional aspects

The South East Asian arboreal Formicine Echinopla melanarctos, as well as some other members of this genus possess a cuticular structure unique in ants, the pedestal hairs. In E. melanarctos, about 700 pedestal hairs are situated on the dorsal and lateral surfaces of the head, the alitrunk, the petiole and the gaster. They are arranged in a polygon-like figuration. On the summit of each of the up to 200-μm high pedestals, a single central hair inserts. This hair (up to 500-μm long) is innervated by a single bipolar mechanosensitive sensory cell. The lumen of each tube-like pedestal contains (1) epithelial cells (2) the sensory cell and the auxiliary cells of the central hair and (3) the long efferent ductules of up to ten isolated bicellular glandular units. Each glandular unit is composed of a secretory glandular cell and a duct cell, all of which are located at the base of a pedestal. The cytoplasm of a glandular cell contains a well-developed end apparatus and is characterised by stacks of smooth and granular endoplasmic reticulum, numerous polyribosomes, a lot of mitochondria and some up to 5-μm large secretory vesicles. The secretion of the gland cells is released on the apex of the pedestal wall via small pores. Approximately 30 μm below their summit, some pedestals possess additionally (up to six) mechanosensitive hairs that are arranged ray-like. We suppose that the pedestal hairs are important in nest-space protection and find that only in ants with high pedestals on the head (Echinopla melanarctos and Echinopla pallipes), the compound eyes are stalked thus overtopping the pedestals.     

ULTRASTRUCTURAL EVIDENCE FOR SECRETORY PHASES IN THE LIFE HISTORY OF STIGMATIC HAIRS OF COTTON: A DRY TYPE STIGMA

Stigmatic hairs of the cotton flower were studied through their developmental stages up to anthesis. Stigmatic hairs invariably develop from a densely straining band of epidermal cells opposite the transmitting tissue cells. At anthesis, these are single cell structures measuring up to 300 μm long. At the 5-mm stage of stylar length (7–10 days before anthesis), some stigmatic hair cells begin to accumulate an osmiophilic substance between the plasmalemma and the cell wall, possibly synthesized in the endoplasmic reticulum. This material is apparently never secreted outside the cell wall. Immediately following this secretory phase in some stigmatic hair cells a second secretory phase starts. A dense osmiophilic substance, different in appearance from the previous phase, accumulates in the vacuoles of each hair cell. Concomitantly, dimorphism develops in the cytoplasmic densities of stigmatic hair. Some stigmatic hair cytoplasm appears very dense and shows signs of degeneration while other cytoplasm appears normal. A third secretory phase, which begins at anthesis, occurs in the normal hair cells. This phase is characterized by enhanced activity in the cytoplasm of the endoplasmic reticulum and Golgi apparatus. Large vesicles containing granular material are seen fusing with the plasmalemma. Coincident with this activity there is dissolution of the middle layers of the cell wall and the cuticle is ruptured at various points. The dense osmiophilic substance that had accumulated in the vacuole breaks down into fine granular material. Significance of these changes is discussed in relation to the pollen germination mechanism on the dry type stigma of cotton.     

Influence of Hairiness ofGerbera jamesoniiLeaves on the Searching Efficiency of the ParasitoidEncarsia formosa

To characterize the relationship between the leaf surface ofGerbera jamesoniiHooker cultivars and the searching behavior of the parasitoidEncarsia formosaGahan on a leaf, the trichome density and shape were described, and the walking behavior was quantified. Leaf hair density varied from 80 to more than 1000 trichomes/cm²and the hair shape varied from single erect trichomes to tapestries of entangled trichomes above the leaf surface. The walking activity, speed, and pattern of the parasitoid were tested on leaves with different hair density and shape. In spite of the large differences in leaf surface structure, the walking activity was similar and around 75% on most cultivars. The walking speed was between 0.2 and 0.3 mm/s on all cultivars and was not significantly different from the speed on tomato. On hairless sweet pepper leaves the walking speed is much higher (0.73 mm/s), so the hairs onGerberaleaves do hamper parasitoid females and result in a strong reduction of the walking speed, but the variation in hair density and shape onGerberadoes not lead to differences in walking activity, pattern, and speed. It appears that the hairs ofGerberaare less of an obstacle forE. formosathan the stiff, large hairs occurring in a high density on cucumber, where the walking speed of the parasitoid is less than 0.2 mm/s. A rise of temperature of 5°C gave a significantly higher walking speed (0.39 mm/s) ofE. formosafemales on all cultivars tested. The relative straightness of the walking track was high and the same on all cultivars. Consequences of the results of the searching behavior ofE. formosaonG. jamesoniicultivars are discussed with respect to other host plants. As parasitoid walking speeds are the same onGerberaand tomato, and whitefly reproduction is also similar on these two host plants, we conclude that biological control of whiteflies onGerberais a realistic option. This conclusion is confirmed by the strong increase of commercial biological control onGerberawhich occurred during the past 5 years.     

Hair morphology inPlantago sect.Coronopus (Plantaginaceae)

The presence of bottle-like hairs has been described in four hitherto uninvestigated taxa ofPlantago sect.Coronopus sensuDietrich. This hair type is a feature characteristic of sect.Coronopus.     

Multiple cyclic nucleotide‐gated channels coordinate calcium oscillations and polar growth of root hairs

Calcium gradients underlie polarization in eukaryotic cells. In plants, a tip‐focused Ca²⁺‐gradient is fundamental for rapid and unidirectional cell expansion during epidermal root hair development. Here we report that three members of the cyclic nucleotide‐gated channel family are required to maintain cytosolic Ca²⁺ oscillations and the normal growth of root hairs. CNGC6, CNGC9 and CNGC14 were expressed in root hairs, with CNGC9 displaying the highest root hair specificity. In individual channel mutants, morphological defects including root hair swelling and branching, as well as bursting, were observed. The developmental phenotypes were amplified in the three cngc double mutant combinations. Finally, cngc6/9/14 triple mutants only developed bulging trichoblasts and could not form normal root hair protrusions because they burst after the transition to the rapid growth phase. Prior to developmental defects, single and double mutants showed increasingly disturbed patterns of Ca²⁺ oscillations. We conclude that CNGC6, CNGC9 and CNGC14 fulfill partially but not fully redundant functions in generating and maintaining tip‐focused Ca²⁺ oscillations, which are fundamental for proper root hair growth and polarity. Furthermore, the results suggest that these calmodulin‐binding and Ca²⁺‐permeable channels organize a robust tip‐focused oscillatory calcium gradient, which is not essential for root hair initiation but is required to control the integrity of the root hair after the transition to the rapid growth phase. Our findings also show that root hairs possess a large ability to compensate calcium‐signaling defects, and add new players to the regulatory network, which coordinates cell wall properties and cell expansion during polar root hair growth.     

Hair,encoding a single C2H2 zinc‐finger protein,regulates multicellular trichome formation in tomato

Trichomes originate from the epidermal cells of nearly all terrestrial plants, which are specialized unicellular or multicellular structures. Although the molecular mechanism regulating unicellular trichome formation has been extensively characterized, most of the genes essential for multicellular trichome formation remain unknown. In this study, we identified an associated locus on the long arm of chromosome 10 using a genome‐wide association study (GWAS) on type‐I trichomes of 180 diverse Solanum lycopersicum (tomato) accessions. Using map‐based cloning we then cloned the key gene controlling the initiation of this type of trichome, named Hair (H), which encodes a single C2H2 zinc‐finger protein. Transgenic experiments showed that hair‐absent phenotype is caused by the deletion of the entire coding region of H. We identified three alleles of H containing several missense mutations and a nucleotide deletion, which result in amino acid substitutions and a reading frame shift, respectively. In addition, knockdown of H or Woolly (Wo) represses the formation of type‐I trichomes, suggesting that both regulators may function as a heterodimer. Direct protein–protein interaction between them was further detected through pull‐down and yeast two‐hybrid assays. In addition, ectopic expression of H in Nicotiana tabacum (tobacco) and expression of its homologs from Capsicum annuum (pepper) and tobacco in tomato can trigger trichome formation. Taken together, these findings suggest that the H gene may be functionally conserved in multicellular trichome formation in Solanaceae species.