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.     

Trichoderma species form endophytic associations within Theobroma cacao trichomes

Trichoderma species are usually considered soil organisms that colonize plant roots, sometimes forming a symbiotic relationship. Recent studies demonstrate that Trichoderma species are also capable of colonizing the above ground tissues of Theobroma cacao (cacao) in what has been characterized as an endophytic relationship. Trichoderma species can be re-isolated from surface sterilized cacao stem tissue, including the bark and xylem, the apical meristem, and to a lesser degree from leaves. SEM analysis of cacao stems colonized by strains of four Trichoderma species (Trichoderma ovalisporum-DIS 70a, Trichoderma hamatum-DIS 219b, Trichoderma koningiopsis-DIS 172ai, or Trichoderma harzianum-DIS 219f) showed a preference for surface colonization of glandular trichomes versus non-glandular trichomes. The Trichoderma strains colonized the glandular trichome tips and formed swellings resembling appresoria. Hyphae were observed emerging from the glandular trichomes on surface sterilized stems from cacao seedlings that had been inoculated with each of the four Trichoderma strains. Fungal hyphae were observed under the microscope emerging from the trichomes as soon as 6 h after their isolation from surface sterilized cacao seedling stems. Hyphae were also observed, in some cases, emerging from stalk cells opposite the trichome head. Repeated single trichome/hyphae isolations verified that the emerging hyphae were the Trichoderma strains with which the cacao seedlings had been inoculated. Strains of four Trichoderma species were able to enter glandular trichomes during the colonization of cacao stems where they survived surface sterilization and could be re-isolated. The penetration of cacao trichomes may provide the entry point for Trichoderma species into the cacao stem allowing systemic colonization of this tissue.     

“HAIRY CANOLA” – Arabidopsis GL3 Induces a Dense Covering of Trichomes on Brassica napus Seedlings

Transformation with the Arabidopsis bHLH gene 35S:GLABRA3 (GL3) produced novel B. napus plants with an extremely dense coverage of trichomes on seedling tissues (stems and young leaves). In contrast, trichomes were strongly induced in seedling stems and moderately induced in leaves of a hairy, purple phenotype transformed with a 2.2 kb allele of the maize anthocyanin regulator LEAF COLOUR (Lc), but only weakly induced by BOOSTER (B-Peru), the maize Lc 2.4 kb allele, or the Arabidopsis trichome MYB gene GLABRA1 (GL1). B. napus plants containing only the GL3 transgene had a greater proportion of trichomes on the adaxial leaf surface, whereas all other plant types had a greater proportion on the abaxial surface. Progeny of crosses between GL3⁺ and GL1⁺ plants resulted in trichome densities intermediate between a single-insertion GL3⁺ plant and a double-insertion GL3⁺ plant. None of the transformations stimulated trichomes on Brassica cotyledons or on non-seedling tissues. A small portion of bHLH gene-induced trichomes had a swollen terminal structure. The results suggest that trichome development in B. napus may be regulated differently from Arabidopsis. They also imply that insertion of GL3 into Brassica species under a tissue-specific promoter has strong potential for developing insect-resistant crop plants. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Interactions among the entomopathogenic fungus, Beauveria bassiana (Ascomycota: Hypocreales), the parasitoid, Aphidius matricariae (Hymenoptera: Braconidae), and its host, Myzus persicae (Homoptera: Aphididae)

The effect of the entomopathogenic fungus Beauveria bassiana on the biological characteristics and life table of Aphidius matricariae, a parasitoid of the green peach aphid, Myzus persicae, was studied under laboratory conditions. Aphids were first infected with twice the LC₉₅ of B. bassiana for third-instar M. persicae (2 × 10⁸ conidia/ml). Subsequently, at different intervals they were exposed to 1-day-old mated parasitoid females for 24 h. The number of mummies produced per female and the percentage emergence of the F1 generation differed significantly as a function of the time interval between application of the fungus and exposure to the parasitoid. The interference of B. bassiana on parasitoid development was also studied by first exposing the aphid hosts to the parasitoid for 24 h and subsequently applying B. bassiana. The number of mummies produced by a female A. matricariae varied from 11.8 to 24.8 and was significantly different when the aphids were first exposed to the parasitoids and then treated with B. bassiana 24, 48, 72, and 96 h after exposure. There were no significantly different effects of B. bassiana on net reproductive rate (R₀), mean generation time (T), intrinsic rate (r_m) and the finite rate of increase (λ) of A. matricariae as a result of development in hosts exposed to low or high conidial concentrations (1 × 10², 2 × 10⁸ conidia/ml). The parasitoids developed in infected hosts had lower r_m, λ, T and DT (doubling time) values compared with those that developed in uninfected hosts but no differences were observed in R₀ values. With proper timing, A. matricariae and B. bassiana can be used in combination in the successful biological control of M. persicae.     

Nonglandular leaf trichomes as short-term inducible defense of the grey alder,Alnus incana (L.), against the chrysomelid beetle,Agelastica alni L.

Summary The grey alder compensates leaf area losses due to insect grazing by continuously producing new leaves throughout the vegetative period. Different degrees of defoliation were attained experimentally by a controlled release of the oligophagous beetle Agelastica alni on arbitrarily selected trees from a homogenous population of young alders. The reduction in leaf area per tree significantly influenced the density of leaf trichomes, assessed 10–30 days later, on newly sprouting leaves only. Cross-correlations between leaf area reduction and trichome density were strongest for leaves which completed unfolding 14–21 days after damage. Dualchoice assays suggested a negative influence of trichomes on oviposition rate of A. alni. Removal of trichomes by shaving demonstrated the highly significant effect of trichomes on feeding behavior of adults and larvae in dual-choice assays. The role of the induced increase in trichome density as a possible short-term defense reaction against herbivorous insects is discussed.     

Resource availability and the trichome defenses of tomato plants

We conducted two experiments to determine how resource availability influenced allocation by tomato (Lycopersicon esculentum) to trichomes, and how different patterns of trichome allocation by plants grown in different resource environments might then influence the behavior of tobacco hornworm (Manduca sexta) caterpillars. In the first experiment we used high and low levels of light and water, and then, using scanning electron microscopy, determined trichome densities on the leaves and stems. We sampled leaves and stems at several places throughout the plant to determine whether there were within-plant differences in allocation to trichomes. The results of the first experiment showed that resource availability influenced allocation to trichome growth. Patterns in high and low-light supported both the growth-differentiation balance hypothesis (GDBH) and the carbon-nutrient balance hypothesis (CNBH). However, the GDBH was not supported by differences among water treatments. Contrary, to predictions of the GDBH, plants with intermediate growth did not have the highest trichome densities, and plants with similar growth differed in trichome density. Possible biological and artifactual explanations are discussed. The first experiment also showed that there was within-plant variation in allocation to trichomes, and that plant resource availability may influence within-plant variation in allocation to trichomes. In the second experiment, we grew plants in high and low-light, and then monitored the behavior of tobacco hornworms on the stems of these plants in the laboratory. This experiment demonstrated that the light environment that tomato plants were grown in influenced the resting behavior of caterpillars. Furthermore, it demonstrated that both glandular and non-glandular trichomes impeded caterpillars from searching for food. Overall, this study indicated that plant resource availability can influence allocation to trichome defenses, and that these differences may affect insect herbivores.     

Accumulation and turnover of 2-tridecanone in Tetranychus urticae and its consequences for resistance of wild and cultivated tomatoes

In this study we assessed the dynamic changes of 2-tridecanone in a herbivorous mite (Tetranychus urticae) on tomato (Lycopersicon esculentum, cv. Moneymaker), a plant with methyl ketones in the tetracellular tips of the glandular trichomes (Type VI). We showed that spider mites accumulate 2-tridecanone when foraging on cultivated tomato. Thus, the rate of mite–trichome contact multiplied by the amount of toxin per trichome tip exceeded the relative rate of toxin turnover multiplied by the amount of toxin per mite. The relative rate of toxin turnover was estimated to be 1.1 per day on cucumber, a plant without this toxin. The amount per trichome tip varied from 0.33 ng for middle-leaf trichomes to 1.26 ng for main-stem trichomes. Hence, to achieve a static level of 2-tridecanone equal to 8–17 ng per mite – representing the level we found in mites on middle leaves – the rate of mite–trichome contact should be 26–57 per day. Because methyl ketone apparently accumulates in the spider mites on tomato, the rate of mite–trichome contact is probably higher than that. We expect the accumulation of ketones to occur especially on the stems of cultivated tomato, since this is the area most densely occupied with glandular hairs and because here the hairs have higher levels of the methyl ketones.Using dose–response relationships assessed earlier (Chatzivasileiadis and Sabelis, 1997, 1998), we estimated that the number of mite–trichome contacts causing 50% mortality per day is equal to 88 on a tomato stem, whereas it equals 70 for another strain of spider mites collected from cucumber. On wild tomato, L. hirsutum f. glabratum (PI 134417), just one to two contacts would suffice to cause 50% mortality per day. We suggest that methyl ketones from glandular hairs on tomato are an important mortality factor for spider mites on wild tomato and probably also on cultivated tomato.     

AFLP markers in the detection of Scirpus × mariqueter (CYPERACEAE) hybrid in China

Scirpus × mariqueter Tang & F.T. Wang (CYPERACEAE) is a pioneer plant of the tidal zone of estuaries of East Asia. Amplified fragment length polymorphisms (AFLP) were used to infer the relationship between S. × mariqueter and its putative progenitors, Scirpus planiculmis F. Schmidt and Scirpus triqueter L. The genetic distance between S. × mariqueter and S. triqueter was much higher than that between S. × mariqueter and S. planiculmis, and samples of S. × mariqueter and those of S. planiculmis did not form a monophyletic group, while the S. triqueter group formed a distinct monophyletic group. However, we also detected an apparent clustering of individuals of S. × mariqueter and S. planiculmis separately. Our results disprove the hybrid origin of S. × mariqueter and rather suggest that it is more closely related to S. planiculmis.     

线纹香茶菜叶上腺毛发育的细胞学研究

为进行中药溪黄草基原植物的品种鉴定,采用光镜和电镜对线纹香茶菜(原变种)[Isodon lophanthoides var.lophanthoides]叶上腺毛的发育进行细胞学研究。结果表明,线纹香茶菜具有头状腺毛和盾状腺毛2种类型。头状腺毛无色透明,由1个基细胞、1个柄细胞和1或2个头部分泌细胞构成;盾状腺毛为红色,由1或2个基细胞、1个柄细胞和4~8个分泌细胞构成头部。2种腺毛均由原表皮细胞经两次平周分裂形成,后因柄细胞和头部细胞所处的分化状态不同而形成两类腺毛。2种腺毛超微结构表明,质体、高尔基体和粗面内质网为主要分泌物产生和运输的细胞器。当盾状腺毛成熟时,角质层下间隙充满了分泌物,其分泌物的性质很可能决定了线纹香茶菜腺毛的颜色。     

Trichome morphology provides phylogenetically informative characters for Tremandra, Platytheca and Tetratheca (former Tremandraceae)

Summary Trichomes of Tremandra R.Br. ex DC., Platytheca Steetz and Tetratheca Sm. (Elaeocarpaceae, former Tremandraceae), together with two outgroup species of Elaeocarpus L., are illustrated using scanning electron microscopy, and their distribution on various plant organs is documented. Various trichomes types were identified that relate taxa: simple hairs, stellate hairs, short glandular trichomes, long glandular trichomes, and three forms of tubercules. Both outgroup and ingroup taxa have simple hairs. Stellate hairs are confirmed as unique to Tremandra. Prominent and sculptured multicelled tubercules, some bearing a stout hair, are characteristic of Platytheca. Smaller multicelled tubercules occur in both Platytheca and Tetratheca, except for the Western Australian taxon Te. filiformis Benth. (possibly plesiomorphic). Unicellular tubercules (papilla) characterise two species of Tetratheca. Short glandular trichomes, usually found on the ovary, also occur in both of these genera but not in all species (possibly secondary losses), while long glandular trichomes, usually on stems and leaves, occur only in some groups of Tetratheca. Within Tetratheca, Western Australian taxa that have five-merous flowers fall into three ‘groups’: seven species (together with one from South Australia) that have short glandular trichomes but no long glandular trichomes; six species that have long glandular trichomes but no short glandular trichomes; and four species that have both trichome types. All other species of Tetratheca have four-merous flowers and form two ‘groups’: 12 eastern species (including one from South Australia) that have both short glandular trichomes and long glandular trichomes; 4 western species and six eastern species that lack short glandular trichomes. On the basis of these characters, a phylogenetic hypothesis for the three genera is presented.     

Molecular identification of intestinal microflora in Takifugu niphobles

We investigated the intestinal microflora of coastal fish including Takifugu niphobles using both culture techniques and library cloning. As a result, the numbers of bacteria appeared on agar media were 1.0 × 10⁴ to 1.4 × 10⁹ CFU/g (colony forming units/gram), whereas those of total bacteria stained with 4′,6-diamidino-2-phenylindole were 4.7 × 10¹⁰ to 1.9 × 10¹¹ cells/gram, irrespective of different fish species. In addition, the culture technique showed that the intestinal microflora in all specimens was mainly composed of the genus Vibrio. In contrast, the direct count method showed that spirochaetes with length of 2.5-4.5 μm were present in the intestinal contents of T. niphobles at high densities, whereas such bacteria could not be detected in those of other fish species. Library cloning yielded the sequences of 16S rRNA genes that were divided into seven taxonomic categories of bacteria including Actinobacteria, Bacilli, Clostridia, Gammaproteobacteria, Mollicutes, Spirochaetes and an unclassified bacterial group. These results demonstrate that the molecular diversity of the intestinal bacteria in T. niphobles based on the clone library method reflects the direct observation by fluorescence microscopy to some extent.     

Ontogenetics of QTL: the genetic architecture of trichome density over time in Arabidopsis thaliana

Although much is known about the molecular genetic basis of trichome development in Arabidopsis thaliana, less is known about the underlying genetic basis of continuous variation in a trait known to be of adaptive importance: trichome density. The density of leaf trichomes is known to be a major determinant of herbivore damage in natural populations of A. thaliana and herbivores are a significant selective force on genetic variation for trichome density. A number of developmental changes occur during ontogeny in A. thaliana, including changes in trichome density. I used multiple interval mapping (MIM) analysis to identify QTL responsible for trichome density on both juvenile leaves and adult leaves in replicate, independent trials and asked whether those QTL changed with ontogeny. In both juvenile and adult leaves, I detected a single major QTL on chromosome 2 that explained much of the genetic variance. Although additional QTL were detected, there were no consistent differences in the genetic architecture of trichome density measured on juvenile and adult leaves. The finding of a single QTL of major effect for a trait of known adaptive importance suggests that genes of major effect may play an important role in adaptation.     

Functional response and progeny production of the Madeira mealybug parasitoid, Anagyrus sp. nov. nr. sinope: The effects of host and parasitoid densities

We tested the hypothesis that control of an herbivorous pest would be improved by providing floral resources for adult natural enemies. The herbivore was euonymus scale, Unaspis euonymi (Comstock) (Homoptera: Diaspididae), a serious pest of woody ornamental plants. The experimental landscape consisted of 3 × 3 m plots, each containing a central bed of Euonymus fortunei (Turcz.) that was infested with the scale. Floral resource plants were cultivars of four species that overlapped in bloom periods to provide a continuous supply of floral resources during summer: Trifolium repens L., Euphorbia epithymoides L., Coreopsis verticillata L. var. ‘Moonbeam,’ and Solidago canadensis L. var. ‘Golden Baby.’ Plots contained either low or high densities of all four species, or no resource plants. Densities of euonymus scale were typically lower in plots containing resource plants than in plots without them. Parasitism by Encarsia citrina (Craw.) (Hymenoptera: Aphelinidae) was rarely influenced by the experimental treatments, flower biomass, whole-plant biomass, or scale density, but in some cases was inversely correlated with density of scales within a generation and in the subsequent generation. Parasitism occasionally reduced densities of scales in plots containing resource plants, but this effect apparently was related to vegetative, not floral qualities of plants. A steady increase in parasitism rate over the three-year course of the experiment across the entire landscape was associated with decreasing density of scales, suggesting a numerical response by the parasitoid population. These findings suggest that the parasitoid is capable of effectively controlling euonymus scale in ornamental landscapes where environmental conditions are favorable.     

Glandular Trichomes on Alfalfa Impede Searching Behavior of the Potato Leafhopper Parasitoid

New cultivars of alfalfa, Medicago sativa L., have been released with glandular trichomes for resistance to potato leafhopper, Empoasca fabae (Harris). Yet, the impact of the glandular trichomes on the primary natural enemy of the leafhopper, Anagrus nigriventris Girault, is unknown. We compared the host searching behavior of the egg parasitoid on four alfalfa clones varying in trichome characters. Female wasps were videotaped on Ranger, a susceptible clone with relatively sparse trichomes, B14, a resistant clone with dense but nonglandular trichomes, and FG12 and FG18, two resistant clones with glandular trichomes. Although the number of leafhopper eggs per stem exposed to wasps did not significantly differ among the four clones, the frequency of foraging and total foraging time were less on the two clones with glandular trichomes than on the two clones with nonglandular trichomes. In addition, an analysis of covariance demonstrated that, although the number of ovipositional probes increased with egg density on a stem, the number of probes on stems with glandular trichomes was significantly less than that on stems without glandular trichomes. The allocation of time by wasps among drumming, probing, and grooming behaviors was similar among the clones. Wasps tended to fly off of clones with glandular trichomes more often than off of clones with nonglandular trichomes. This study suggests that cultivars with glandular trichomes may interfere with host searching by A. nigriventris.     

6-Benzylaminopurine treatment induces increased pubescence on wheat leaves

The epidermis of wheat (Triticum aestivum L.) leaves contains trichomes that contribute to resistance to insect pests and drought tolerance. In the present study, we examined the effects of 6-benzylaminopurine (BA) and methyl jasmonate (MeJA) treatment on trichome development on the leaves of wheat cv. Norin 61 seedlings. Without phytohormone treatment, trichomes on the adaxial leaf surface were short (90 μm) and their density was low (3.6 trichomes/mm²). Both BA and MeJA treatments significantly increased the density of trichomes, and there were no significant differences between the phytohormone treatments. BA treatment increased trichome length to five times as long as that in the control, whereas MeJA treatment did not significantly affect trichome length. Since BA treatment concurrently increased the DNA content of the nuclei in trichome cells, endoreduplication of the nuclei is probably involved in trichome enlargement. These results indicate that even wheat cultivars with short trichomes retain the mechanisms for trichome enlargement and stimuli such as BA application can induce increased pubescence on wheat leaves.     

Trichomes: their Diversity,Distribution and Density in Acclimatization of Teak (Tectona grandis L.) Plants Grown in vitro

Diversity, distribution and density of trichomes were studied in relation to other morphological and physiological parameters of in vivo and in vitro teak (Tectona grandis L.) plants with the objective to understand their role in hardening and acclimatization of micropropagated plants. Less diversity and scanty distribution of trichomes of in vitro leaves in comparison to the in vivo ones were correlated with water loss of the former plants leading to low rate of survival after transplantation. The probable cause for absence of certain types of trichome in in vitro plants was also discussed.     

Biological control of brown rot (Moniliana laxa Ehr.) on apricot (Prunus armeniaca L. cv. Hacıhaliloğlu) by Bacillus, Burkholdria, and Pseudomonas application under in vitro and in vivo conditions

In 2002 and 2003, a study was conducted to determine the effect of bacterial strains, Burkholdria OSU 7, Bacillus OSU 142, and Pseudomonas BA 8, on biological control of brown rot disease (Monilinia laxa Ehr.) on apricot cv. Hacıhaliloğlu in Malatya province of Turkey. Apricot orchard at full blooming stage was inoculated with conidial suspension (1 × 10⁶ spores/ml) of M. laxa Ehr. After inoculation, two apricot trees for each application were treated with each of the three biological control agents (Burkholdria gladii OSU 7, Bacillus subtilis OSU 142, and Pseudomonas putida BA 8) by spraying (1 × 10⁹ cfu/ml) on inoculated branches. Disease incidence was evaluated for untreated (control 1) and four different treatment groups including commercial disease management (control 2, positive control: 3% Bourdox in fall, 50% Cupper at pink flower, 30 g/100 l Corus at first blooming, and 300 g/100 l Captan at last blooming stage) and treatments including each of the three bacterial strains (OSU 7, OSU 142, and BA 8). The results showed that disease incidence for negative control (control 1) was 9.94, which was significantly higher than disease incidence for commercial application (2.57%) or bacterial treatments (2.82–5.00%) in the first year. In 2003, the lowest disease incidence observed in OSU 7 treatment (6.80%), while disease incidence rate for positive control and negative control were 9.45% and 28.46%, respectively. This result may suggest that OSU 7 has potential to be used as biopesticide for effective management of brown rot disease on apricot.