Induction Of Trichomes Research Articles

Cultivar-Specific Defense Responses in Wild and Cultivated Squash Induced by Belowground and Aboveground Herbivory

Plant domestication often alters plant traits, including chemical and physical defenses against herbivores. In squash, domestication leads to reduced levels of cucurbitacins and leaf trichomes, influencing interactions with insects. However, the impact of domestication on inducible defenses in squash remains poorly understood. Here, we investigated the chemical and physical defensive traits of wild and domesticated squash (Cucurbita argyrosperma), and compared their responses to belowground and aboveground infestation by the root-feeding larvae and the leaf-chewing adults of the banded cucumber beetle Diabrotica balteata (Coleoptera: Chrysomelidae). Wild populations contained cucurbitacins in roots and cotyledons but not in leaves, whereas domesticated varieties lacked cucurbitacins in all tissues. Belowground infestation by D. balteata larvae did not increase cucurbitacin levels in the roots but triggered the expression of cucurbitacin biosynthetic genes, irrespective of domestication status, although the response varied among different varieties. Conversely, whereas wild squash had more leaf trichomes than domesticated varieties, the induction of leaf trichomes in response to herbivory was greater in domesticated plants. Leaf herbivory varied among varieties but there was a trend of higher leaf damage on wild squash than domesticated varieties. Overall, squash plants responded to both belowground and aboveground herbivory by activating chemical defense-associated gene expression in roots and upregulating their physical defense in leaves, respectively. While domestication suppressed both chemical and physical defenses, our findings suggest that it may enhance inducible defense mechanisms by increasing trichome induction in response to herbivory.

Defoliation by Gypsy Moth (Lepidoptera, Erebidae) Induces Differential Delayed Induction of Trichomes in Two Birch Species.

Outbreaks of the gypsy moth (Lymantria dispar japonica Motschulsky) cause serious defoliation in birch. A single year of defoliation has no significant impact on the trees, whereas continuous defoliation events could be fatal. How birch species avoid serious damage caused by gypsy moth outbreak is yet to be revealed. Trichomes on leaf surfaces of birch trees are an effective antiherbivore defense strategy. We examined a 1-yr delayed induction of glandular (GT) and nonglandular trichomes (NGT) on leaf surfaces caused by stress in white birch (Betula platyphylla Sukaczev var. japonica [Miq.] Hara) and monarch birch (Betula maximowicziana Regel). Saplings were subjected to four treatments in June 2009: herbivory (50% of the leaf area was grazed by gypsy moths), mechanical cutting (50% of the leaf area was cut using scissors), shading (50% light shading with a black sheet), and control (covered with a net to prevent herbivory). Then, the density of GT and NGT on early leaves was determined in April 2010. In both birch species, the density of GT was higher in herbivory than in other treatments. The density of GT due to mechanical cutting was higher than that in the control, but only for white birch. However, the density of NGT was lower after mechanical cutting than in other treatments for white birch. There were no differences in the density of NGT among treatments for monarch birch. These results show that 1-yr delayed induction of GT by herbivory was stronger in white birch than in monarch birch.

Variasi morfologi puparium Bemisa tabaci (Gennadius) (Hemiptera: Aleyrodidae) pada berbagai inang dan ketinggian tempat dari daerah endemik penyakit kuning cabai di Wilayah Sundaland

Bemisia tabaci (Gennadius) dikenal sebagai salah satu kutukebul vektor virus gemini yang bersifat polifag. Identifikasi B. tabaci pada umumnya berdasarkan kantung nimfa instar keempat (puparium) yang memiliki mata berwarna merah. Bentuk nimfa instar keempat tersebut memiliki variasi morfologi. Adanya variasi morfologi ini menyebabkan kesulitan dalam identifikasi tingkat spesies. Penelitian ini bertujuan untuk mempelajari variasi puparium B. tabaci pada tanaman inang di berbagai ketinggian tempat di atas permukaan laut. Sampel puparium B. tabaci diperoleh dari daerah endemik di Wilayah Sundaland penyakit kuning cabai di Sumatera Barat, Jawa Barat, Jawa Tengah, Jawa Timur, Bali, dan Kalimantan Selatan. Sebelum dilakukan analisa morfologi, puparium terlebih dahulu dilakukan uji molekuler dan dinyatakan sebagai B. tabaci. Puparium kemudian dipreparasi dalam preparat mikroskop dan dilakukan identifikasi morfologi pada level spesies. Selanjutnya dilakukan pengamatan ukuran dan bentuk puparium serta pengamatan jumlah seta dorsal dan ukuran seta kauda. Analisis korelasi kanonik digunakan untuk mengetahui faktor yang mempengaruhi variasi puparium. Ada empat kelompok bentuk puparium B. tabaci, yaitu oval, oval dengan 1–2 lekukan, oval dengan >3 lekukan, dan bentuk seperti kerang laut. Terdapat hubungan antara jenis tanaman inang dan variasi morfologi puparium. Trikoma yang ada di permukaan daun tanaman inang menyebabkan terjadinya variasi jumlah seta dorsal yang memanjang, variasi bentuk, dan ukuran puparium. Variasi puparium B. tabaci lebih dipengaruhi oleh jenis tanaman inang daripada ketinggian tempat di atas permukaan laut.

Genome-wide characterization of NtHD-ZIP IV: different roles in abiotic stress response and glandular Trichome induction

BackgroundThe plant-specific homeodomain-leucine zipper class IV (HD-ZIP IV) gene family has been involved in the regulation of epidermal development.ResultsFifteen genes coding for HD-ZIP IV proteins were identified (NtHD-ZIP-IV-1 to NtHD-ZIP-IV-15) based on the genome of N. tabacum. Four major domains (HD, ZIP, SAD and START) were present in these proteins. Tissue expression pattern analysis indicated that NtHD-ZIP-IV-1, − 2, − 3, − 10, and − 12 may be associated with trichome development; NtHD-ZIP-IV-8 was expressed only in cotyledons; NtHD-ZIP-IV-9 only in the leaf and stem epidermis; NtHD-ZIP-IV-11 only in leaves; and NtHD-ZIP-IV-15 only in the root and stem epidermis. We found that jasmonates may induce the generation of glandular trichomes, and that NtHD-ZIP-IV-1, − 2, − 5, and − 7 were response to MeJA treatment. Dynamic expression under abiotic stress and after application of phytohormones indicated that most NtHD-ZIP IV genes were induced by heat, cold, salt and drought. Furthermore, most of these genes were induced by gibberellic acid, 6-benzylaminopurine, and salicylic acid, but were inhibited by abscisic acid. NtHD-ZIP IV genes were sensitive to heat, but insensitive to osmotic stress.ConclusionNtHD-ZIP IV genes are implicated in a complex regulatory gene network controlling epidermal development and abiotic stress responses. The present study provides evidence to elucidate the gene functions of NtHD-ZIP IVs during epidermal development and stress response.

Tobacco transcription repressors NtJAZ: Potential involvement in abiotic stress response and glandular trichome induction

Members of the Jasmonate ZIM domain (JAZ) proteins act as transcriptional repressors in the jasmonate (JA) hormonal response. To characterize the potential roles of JAZ gene family in plant development and abiotic stress response, fifteen JAZs were identified based on the genome of Nicotiana tabacum. Structural analysis confirmed the presence of single Jas and TIFY motif. Tissue expression pattern analysis indicated that NtJAZ-2, -3, -5, and -10 were highly expressed in roots and NtJAZ-11 was expressed only in the cotyledons. The transcript level of NtJAZ-3, -5, -9, and -10 in the stem epidermis was higher than that in the stem without epidermis. Dynamic expression of NtJAZs exposed to abiotic stress and phytohormone indicated that the expression of most NtJAZs was activated by salicylic acid, methyl jasmonate, gibberellic acid, cold, salt, and heat stresses. With abscisic acid treatment, NtJAZ-1, -2, and -3 were not activated; NtJAZ-4, -5, and -6 were up-regulated; and the remaining NtJAZ genes were inhibited. With drought stress, the expression of NtJAZ-1, -2, -3, -4, -5, -6, -7, and -8 was up-regulated, whereas the transcript of the remaining genes was inhibited. Moreover, high concentration MeJA (more than 1 mM MeJA) had an effect on secreting trichome induction, but inhabited the plant growth. Nine NtJAZs may play important role in secreting trichome induction. These results indicate that the JAZ proteins are convergence points for various phytohormone signal networks, which are involved in abiotic stress responses.

12-Hydroxy-Jasmonoyl-l-Isoleucine Is an Active Jasmonate That Signals through CORONATINE INSENSITIVE 1 and Contributes to the Wound Response in Arabidopsis.

12-hydroxy-jasmonoyl-isoleucine (12OH-JA-Ile) is a metabolite in the catabolic pathway of the plant hormone jasmonate, and is synthesized by the cytochrome P450 subclade 94 enzymes. Contrary to the well-established function of jasmonoyl-isoleucine (JA-Ile) as the endogenous bioactive form of jasmonate, the function of 12OH-JA-Ile is unclear. Here, the potential role of 12OH-JA-Ile in jasmonate signaling and wound response was investigated. Exogenous application of 12OH-JA-Ile mimicked several JA-Ile effects including marker gene expression, anthocyanin accumulation and trichome induction in Arabidopsis thaliana. Genome-wide transcriptomics and untargeted metabolite analyses showed large overlaps between those affected by 12OH-JA-Ile and JA-Ile. 12OH-JA-Ile signaling was blocked by mutation in CORONATINE INSENSITIVE 1. Increased anthocyanin accumulation by 12OH-JA-Ile was additionally observed in tomato and sorghum, and was disrupted by the COI1 defect in tomato jai1 mutant. In silico ligand docking predicted that 12OH-JA-Ile can maintain many of the key interactions with COI1-JAZ1 residues identified earlier by crystal structure studies using JA-Ile as ligand. Genetic alternation of jasmonate metabolic pathways in Arabidopsis to deplete both JA-Ile and 12OH-JA-Ile displayed enhanced jasmonate deficient wound phenotypes and was more susceptible to insect herbivory than that depleted in only JA-Ile. Conversely, mutants overaccumulating 12OH-JA-Ile showed intensified wound responses compared with wild type with similar JA-Ile content. These data are indicative of 12OH-JA-Ile functioning as an active jasmonate signal and contributing to wound and defense response in higher plants.

Salt tolerance traits in Deschampsia antarctica Desv.

AbstractDeschampsia antarctica Desv. (Poaceae) grows in coastal habitats in the Maritime Antarctic where it is often exposed to sea spray. Salt crystals have been observed on the surface of leaves in plants treated with high NaCl. We investigated if D. antarctica is a salt tolerant species that allows sodium ions to diffuse into the root where a salt overly sensitive (SOS) system extrudes Na+ from root cells and facilitates its movement through the xylem up to the leaves. Leaf epidermis, physiological parameters and sodium transporters in D. antarctica plants exposed to NaCl were studied over 21 days. Epidermal scanning electron microscopy showed trichome induction in the leaves of salt treated plants. In addition, salt treated plants showed increased sodium and proline levels with a concomitant increased expression of SOS genes (1 and 3). These results indicate that Na+ is taken up by the roots of D. antarctica and transported to the leaves. The sodium flux may be controlled by SOS1 activity. Up-regulation of the SOS1 gene may be involved in the increased sodium levels observed in the leaves of salt treated plants. Trichomes may also be involved in sodium exudation through the leaves under saline conditions.

The Genetics of Phenotypic Plasticity in Plant Defense: Trichome Production inMimulus guttatus

Insect herbivory is a major driving force of plant evolution. Phenotypic plasticity and developmental variation provide a means for plants to cope with variable herbivory. We characterized the genetics of developmental variation and phenotypic plasticity in trichome density, a putative defensive trait of Mimulus guttatus (yellow monkeyflower). Our results are evaluated in relation to the optimal defense theory, which provides testable predictions for plastic and developmental patterns in defense traits. We found that both developmental stage and simulated insect damage affected trichome production, but in different ways. Plants were more likely to produce at least some trichomes on later leaves than on earlier leaves, regardless of damage. Damage did not affect the average probability of producing trichomes, but it did increase the density of hairs on trichome-positive plants. We mapped trichome quantitative trait loci (QTL) by selectively genotyping a large panel of recombinant inbred lines derived from two highly divergent populations. Several highly pleiotropic QTL influenced multiple aspects of the trichome phenotype (constitutive, developmental, and/or plastic responses). Only one of the QTL influenced trichome induction following damage. In a result that is consistent with a central prediction of optimal defense theory, the high allele at this location was from the ancestral population with low constitutive trichome production.

Jasmonic acid control of GLABRA3 links inducible defense and trichome patterning inArabidopsis

Once attacked by herbivores, plants regenerate new leaves with increased trichome density as an inducible defense. Trichomes are specified from neighboring epidermal cells through local cell-cell interactions in the leaf primordia. However, the molecular mechanism of how herbivore-induced damage at older leaves remodels the pattern of trichome fate specification at newly forming leaves is largely unknown. In this study, we show that mutations in either the biosynthetic or signaling pathway of jasmonates (JAs), long-distance wound signals, abolish the wound-induced formation of trichomes. To identify the factors linking JA signaling to trichome fate specification, we isolated a novel class of mutants, unarmed (urm), which lack trichome induction but show otherwise normal responses to JAs. URM9 encodes an Importin beta family protein, and URM23 is identical to TRANSPARENT TESTA GLABRA1 (TTG1), the product of which interacts with the bHLH transcription factor GLABRA3 (GL3). Loss of either URM9 or URM23 disrupts the subnuclear localization of GL3, thus implicating GL3 in trichome induction. The expression of GL3 was enhanced by JA treatment prior to trichome initiation. Genetic analysis of multiple trichome mutants shows that GL3, in concert with the R2R3-Myb transcription factor GLABRA1 (GL1), promotes trichome fate in response to JA in a dosage-dependent manner. These results indicate that GL3 is a key transcription factor of wound-induced trichome formation acting downstream of JA signaling in Arabidopsis.

Methyl Jasmonate Application Induces Increased Densities of Glandular Trichomes on Tomato, Lycopersicon esculentum

This study was designed to address whether applications of methyl jasmonate (MJ) or Benzothiadiazole (BTH) to cultivated tomato, Lycopersicon esculentum, induced elevated densities of defense-related glandular trichomes on new leaves. Four-leaf tomato plants were sprayed with MJ, BTH, or control solutions, and the density of type VI glandular trichomes on new leaves was subsequently determined at 3, 7, 14, 21, and 28 d. At 7, 14, and 21 d, the density of type VI glandular trichomes on new leaves was significantly higher on MJ-treated plants than on BTH- or control-treated plants. At 7 and 14 d after treatment, the mean density of glandular trichomes on new leaves of MJ-treated plants was ninefold higher than on leaves of control-treated plants. We observed entrapment of immature western flower thrips in trichomes on MJ-treated plants at higher rates than on BTH or control plants. Studies to evaluate potential trade-offs between reductions in pest populations by increased trichome density and possible negative impacts of trichome induction on biological control agents are needed.

Interactive effects of jasmonic acid, salicylic acid, and gibberellin on induction of trichomes in Arabidopsis.

Leaf trichomes protect plants from attack by insect herbivores and are often induced following damage. Hormonal regulation of this plant induction response has not been previously studied. In a series of experiments, we addressed the effects of artificial damage, jasmonic acid, salicylic acid, and gibberellin on induction of trichomes in Arabidopsis. Artificial damage and jasmonic acid caused significant increases in trichome production of leaves. The jar1-1 mutant exhibited normal trichome induction following treatment with jasmonic acid, suggesting that adenylation of jasmonic acid is not necessary. Salicylic acid had a negative effect on trichome production and consistently reduced the effect of jasmonic acid, suggesting negative cross-talk between the jasmonate and salicylate-dependent defense pathways. Interestingly, the effect of salicylic acid persisted in the nim1-1 mutant, suggesting that the Npr1/Nim1 gene is not downstream of salicylic acid in the negative regulation of trichome production. Last, we found that gibberellin and jasmonic acid had a synergistic effect on the induction of trichomes, suggesting important interactions between these two compounds.

Differential induction of trichomes by three herbivores of black mustard

Specificity of plant induction responses may be important to the interactions between mustards and insect herbivores. This study compared the effects of the cabbage white butterfly, Pieris rapae (L.), cabbage looper, Trichoplusia ni (Hubner), and the mustard flea beetle, Phyllotreta cruciferae (Goeze) on induction of leaf trichome density, sinigrin concentration, and nitrogen concentration in black mustard, Brassica nigra (L.) Koch. Plants were damaged for 12h at the four-leaf stage, with effort made to standardize the damage applied. Induction responses were measured on the fifth, seventh, ninth, and 11th leaves counted from the cotyledons. Seventh leaves of plants damaged by P. rapae had 76% more trichomes per unit area than controls, whereas equivalent leaves of plants damaged by the other two herbivores exhibited no response. Ninth leaves of plants damaged by T. ni had 113% more trichomes per unit area than controls, whereas equivalent leaves of plants damaged by the other two herbivores exhibited no response. Trichome densities of fifth and 11th leaves did not respond to treatments. Leaf sinigrin and nitrogen concentrations were not affected by the damage treatments. Differential plant trichome response to P. rapae and T. ni may have been due to differences in location of feeding during the damage treatment. Other cues, such as salivary components, may also have differed between the two herbivores. This study is one of the first to document differential effects of two herbivores from the same guild on induction of morphological resistance.

Tissue layer and organ specificity of trichome formation are regulated by GLABRA1 and TRIPTYCHON in Arabidopsis

In animal development, cellular diversity is generated within tissues which in turn are derived from germ layers. Similar to the germ layers in animals, plants establish three distinct tissue layers early in development which each give rise to a distinct set of cell types. To investigate the role of tissue-layer-specific cues in generating plant cellular diversity we studied the spatial regulation of an epidermal cell type, trichomes (hairs), by the two genes, GLABRA1 (GL1) and TRIPTYCHON (TRY). Ubiquitous expression of the positive regulator GL1 in the absence of the negative regulator TRY leads to ectopic trichome formation not only on additional organs but also in subepidermal tissue layers. Trichomes in inner tissue layers can differentiate the same morphology and show a spacing pattern comparable to trichomes in the epidermis. This clearly shows that cell type specification takes place downstream of tissue-specific cues. We propose a model of how the tissue and organ specificity of trichome induction is regulated in normal development.