Morphogenesis of galls induced by Baccharopelma dracunculifoliae (Hemiptera: Psyllidae) on Baccharis dracunculifolia (Asteraceae) leaves.

The commonest insect gall on Baccharis dracunculifolia (Asteraceae) leaves is induced by Baccharopelma dracunculifoliae (Hemiptera, Psyllidae). The gall-inducing insect attacks young leaves in both the unfolded and the fully expanded stages. Four developmental phases were observed in this type of gall: 1) A folding phase, during which the leaf lamina folded upward alongside the midrib and the edges of the upper portion of the leaf approached each other, forming a longitudinal slit. A single chamber was formed on the adaxial surface of the leaf; 2) A swelling phase, in which the folded leaf tissues thickened and the edges of the leaf drew closer together, narrowing the slit. In this phase the gall matured, turning succulent, fusiform and pale green. The single nymphal chamber was lined with white wax and was able to house from one to several nymphs; 3) A dehiscence phase, characterized by the opening of the slit to release inducers; and 4) A senescence phase, when the gall turned dark and dry. The dermal system of the mature gall was composed of a single-layered epidermis. The mesophyll was swollen, and the swelling was due mainly to hyperplasia of the parenchyma. The vascular tissues along the midrib vein were conspicuous and the perivascular fibers resembled parenchymal cells. The hypertrophied secretory cavities contained low lipophylic content. This gall does not form nutritive tissue, but salivary sheaths left by the inducers were observed near the parenchyma, vascular bundles and secretory cavities. This study complements our current knowledge of gall biology and sheds further light on the plasticity of plant tissues stimulated by biotic factors.     

GENOTYPIC VARIATION AND INTEGRATION IN HISTOLOGICAL FEATURES OF THE GOLDENROD BALL GALL

The herb Solidago altissima (Compositae) forms a gall when attacked by the fly Eurosta solidaginis (Diptera: Tephritidae). We performed an experiment to determine how histological features of gall morphology contribute to previously observed genetic variation in gall diameter. Galls induced on 10 replicated plant clones were sectioned, and their internal tissues and cells measured. Four concentric tissue zones were identified, including a cytoplasmically rich tissue lining the insect's central chamber, and a cytoplasmically poor cortex. As expected, outer gall diameter differed among clones, with a broad sense heritability of 0.29. However, clonal differences for component tissue thicknesses were found for only two of the zones; cortex thickness showed the greater genotypic variation, with a broad sense heritability of 0.30. Path analysis was used to examine the contribution of the zone thicknesses, vascular supply and cell sizes to outer gall diameter at the phenotypic, genotypic and environmental-developmental levels. These showed that cortex thickness was the single greatest determinant of gall size. At the phenotypic level, the single most important determinant of cortex thickness was the number of cortical cells, whereas the amount of cortical vascularization was second in importance. But, at the genotypic level of analysis, vascularization was the single most important determinant of genotypic differences in gall size. The larger the gall, the greater the protection to the inducing insect. Since the bulk of the gall consists of large, cytoplasmically poor cells, the insect gains protection while imposing a small cost on the plant.     

Vascular implications of Dasineura sp. galls' establishment on Peumus boldus stems

• Some chewing larvae are capable of inducing galls in the host vascular cylinder, e.g. Dasineura sp. (Cecidomyiidae) on Peumus boldus stems. Due to the medicinal and economic importance of P. boldus, the anatomical and functional implications of establishment of Dasineura sp. on P. boldus stems were investigated. We asked if establishment of Dasineura sp. in P. boldus stems induces abnormalities at the cellular and organizational level of the vascular system that increase during gall development in favour of the hydric status of the gall.
• Anatomical alterations induced in the stems during gall development were determined. Cytohistometric analyses in mature galls were compared to non-galled stems, and water potential and leaf area of non-galled stems were compared with galled stems.
• Dasineura sp. establishes in the vascular cambium, leading to delignification and rupture of xylem cells, inhibiting formation of phloem and perivascular sclerenchyma. Gall diameter increases together with larval feeding activity, producing a large larval chamber and numerous layers of nutritive tissue, vascular parenchyma, and sclerenchyma. These anatomical alterations do not affect the leaf area of galled stems but favour increased water flow towards these stems.
• The anatomical alterations induced by Dasineura sp. in P. boldus stems guarantee water and nutrient supply to the gall and larva. After the inducer exits stems, some host branches no longer have vascular connections with the plant body.

     

ANOMALOUS GROWTH AND VEGETATIVE ANATOMY OF SIMMONDSIA CHINENSIS

The anatomy of the stem, root, and leaf of Simmondsia chinensis (Link) Schneider was investigated, as well as the mode of tissue formation in the stem. Perivascular tissue is present as part of the primary body; outermost cell layers of this tissue mature as a fibrous sheath. The first short-lived extrafascicular cambium is generated within the remaining parenchymatous perivascular tissue. Successive independent extrafascicular cambia, organized as complete rings or large arcs, arise within peripheral conjunctive parenchyma produced by previous cambia. Extrafascicular cambia produce secondary xylem centripetally and conjunctive tissue bands and strands of secondary phloem centrifugally. Conjunctive tissue initials produce raylike structures of conjunctive tissue; true vascular rays are absent. The phellogen is actually a region of transition where the peripheral conjunctive parenchyma of previous extrafascicular cambia undergoes further cellular subdivision; a true phellogen is lacking. Xylem bands do not represent annual or seasonal growth increments, and secondary growth in Simmondsia is an unequivocal example of the “concentric” anomaly.     

Cynipid galls: insect-induced modifications of plant development create novel plant organs

Cynipid gall formation is achieved by an insect–plant interaction whereby cynipid gallwasps redirect host‐plant development to form novel structures to protect and nourish the developing larvae. Work was carried out to investigate the molecular mechanisms involved in this interaction, and extend the understanding of plant tissue development. Cytological changes of the inner‐gall tissue throughout the development of several gall species was investigated and the developmental stages of gall formation defined, to reveal two different patterns of development followed by the galls tested. Fluorescent in situ hybridization demonstrated many of the inner‐gall cells to be polytenized. Comparisons between inner‐gall and non‐gall tissue protein signatures by Schönrogge et al. (Plant, Cell and Environment 23, 215–222, 2000) have demonstrated the variation between gall and non‐gall protein signatures, and identified a number of inner‐gall proteins. Further analysis of one of these inner‐gall proteins involved in lipid synthesis, putative biotin carboxyl carrier protein (BCCP), revealed differential expression throughout development, and showed this expression to be concentrated in the inner‐gall tissue in all the gall species tested.     

Evidence for long-distance, chemical gall induction by an insect

Abstract We report that a chemical stimulus from a herbivore, a galling insect, changes plant morphology and physiology to benefit the herbivore. Previous studies could not determine whether insect galls are induced by mechanical or chemical stimuli because feeding and oviposition both occurred at the site of gall formation. We report that the mouthparts of a spruce‐galling insect, Adelges cooleyi, were inserted in stem phloem cells far from induced galls, that tissues between mouthparts and galls appeared normal, and that the ability to initiate galls was inversely correlated with distance from buds (potential gall sites). Thus the effects of chemical stimuli were unambiguously separated from any mechanical influence of probing stylets or ovipositors. Our results strongly suggest that galls were induced by a chemical stimulus transported to buds via vascular tissue and that its efficacy was dose‐dependent.     

爱玉子上幼期爱玉子榕小蜂与虫瘿的协同发育

[目的]特种果树爱玉子Ficus pumila var.awkeotsang依赖榕小蜂传粉方能结实.本研究旨在了解爱玉子榕小蜂Wiebesia sp.nr.pumilae及其虫瘿发育动态,为榕小蜂发育生物学研究奠定基础.[方法]通过人工放蜂-标记-定期采样-显微镜和电镜观测的方法,观察爱玉子榕小蜂-虫瘿协同发育过程以及...     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

Manipulation of host plant cells and tissues by gall-inducing insects and adaptive strategies used by different feeding guilds

Biologists who study insect-induced plant galls are faced with the overwhelming diversity of plant forms and insect species. A challenge is to find common themes amidst this diversity. We discuss common themes that have emerged from our cytological and histochemical studies of diverse neotropical insect-induced galls. Gall initiation begins with recognition of reactive plant tissues by gall inducers, with subsequent feeding and/or oviposition triggering a cascade of events. Besides, to induce the gall structure insects have to synchronize their life cycle with plant host phenology. We predict that reactive oxygen species (ROS) play a role in gall induction, development and histochemical gradient formation. Controlled levels of ROS mediate the accumulation of (poly)phenols, and phytohormones (such as auxin) at gall sites, which contributes to the new cell developmental pathways and biochemical alterations that lead to gall formation. The classical idea of an insect-induced gall is a chamber lined with a nutritive tissue that is occupied by an insect that directly harvests nutrients from nutritive cells via its mouthparts, which function mechanically and/or as a delivery system for salivary secretions. By studying diverse gall-inducing insects we have discovered that insects with needle-like sucking mouthparts may also induce a nutritive tissue, whose nutrients are indirectly harvested as the gall-inducing insects feeds on adjacent vascular tissues. Activity of carbohydrate-related enzymes across diverse galls corroborates this hypothesis. Our research points to the importance of cytological and histochemical studies for elucidating mechanisms of induced susceptibility and induced resistance.     

INITIAL TWIG INJURY TO HIGH ELEVATION PICEA RUBENS IN EASTERN NORTH AMERICA CAUSED BY INADEQUATE PERIDERM FORMATION

Apical dieback is the predominant injury symptomatology associated with growth declines of high elevation Picea rubens Sarg, trees in North America. Histological observations of uninjured tissues and of initial injury to tissues were made to understand the mechanism of injury to this species. Observations were made of hundreds of grab samples of apparently uninjured tissues and of uninjured twigs from trees growing on mountains of the Adirondack Mt., NY; Mount Mansfield, VT; Mt. Mitchell, NC; and Clingman's Dome, TN, from March 1985 to April 1986. In the normal growth pattern of red spruce, three buds elongate from each twig terminus during spring. These buds expand into shoot increments during the growing season and three new buds will form at the tip of each of the three elongated increments. The timing of developmental events varied markedly among buds of individual trees and among trees. Bud break occurred between mid-June to the end of July. Most shoot elongation was completed and periderm formation began near the end of August or in early September. A phellogen, one cell thick, formed and a phellem layer developed from phellogen derivatives during autumn 1985. Many twig samples taken in October and November had produced only one, or at most two, phellem layers external to the phellogen during the relatively short growing season. In some samples, three or more phellem layers were present between November 1985 and March 1986. In some cases a distinct phellem was not developed at all. Usually a phelloderm one cell thick developed in autumn of the first year. Tissue necrosis occurred in twigs during their first overwintering period. Injured twigs with necrotic tissues had only one or two continuous or discontinuous phellem layers. In samples that exhibited initial injury, necrotic tissues consisted mostly of cortical cells and phloem subjacent to this meager periderm. Frequently, necrotic tissues developed initially near the bases of needles and at branch “nodes” (transition zone tissues with older twigs). In contrast, twigs of healthy appearance had two or more continuous phellem layers external to the phellogen.     

Phytohormones in Japanese Mugwort Gall Induction by a Gall-Inducing Gall Midge

A variety of insect species induce galls on host plants. Liquid chromatographic/tandem mass spectrometric analyses showed that a gall midge (Rhopalomyia yomogicola) that induces galls on Artemisia princeps contained high levels of indole-3-acetic acid and cytokinins. The gall midge larvae also synthesized indole-3-acetic acid from tryptophan. Close observation of gall tissue sections indicated that the larval chamber was surrounded by layers of cells having secondary cell walls with extensive lignin deposition, except for the part of the gall that constituted the feeding nutritive tissue which was composed of small cells negatively stained for lignin. The differences between these two types of tissue were confirmed by an expression analysis of the genes involved in the synthesis of the secondary cell wall. Phytohormones may have functioned in maintaining the feeding part of the gall as fresh nutritive tissue. Together with the results in our previous study, those presented here suggest the importance of phytohormones in gall induction.     

Histological Comparisons of Fergusobia/Fergusonina-Induced Galls on Different Myrtaceous Hosts

The putative mutualism between different host-specific Fergusobia nematodes and Fergusonina flies is manifested in a variety of gall types involving shoot or inflorescence buds, individual flower buds, stems, or young leaves in the plant family Myrtaceae. Different types of galls in the early-to-middle stages of development, with host-specific species of Fergusobia/Fergusonina, were collected from Australian members of the subfamily Leptospermoideae (six species of Eucalyptus, two species of Corymbia, and seven species of broad-leaved Melaleuca). Galls were sectioned and histologically examined to assess morphological changes induced by nematode/fly mutualism. The different gall forms were characterized into four broad categories: (i) individual flower bud, (ii) terminal and axial bud, (iii) ''basal rosette'' stem, and (iv) flat leaf. Gall morphology in all four types appeared to result from species-specific selection of the oviposition site and timing and number of eggs deposited in a particular plant host. In all cases, early parasitism by Fergusobia/Fergusonina involved several layers of uninucleate, hypertrophied cells lining the lumen of each locule (gall chamber where each fly larva and accompanying nematodes develop). Hypertrophied cells in galls were larger than normal epidermal cells, and each had an enlarged nucleus, nucleolus, and granular cytoplasm that resembled shoot bud gall cells induced by nematodes in the Anguinidae.     

Environmental thermal levels affect the phenological relationships between the chestnut gall wasp and its parasitoids

Studies of thermal level‐related asynchrony in a host–parasitoid relationship are necessary to understand the effects of climate change on new host–parasitoid interactions. In the Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae) and its Chalcidoidea parasitoids, phenological synchrony is assumed to be weather‐dependent in a new area of expansion. To evaluate the effects of environmental thermal regimes on the host, a phenology model for different cynipid stages (larvae, pupae, adults, and adult emergence) and a host–parasitoid phenological estimator are developed in three chestnut fields during two successive growth seasons and subsequently validated in areas with chestnut fields at two different altitudes. Comparisons of the timings of the juvenile and adult stages with those of the parasitoid complex demonstrate that the shortest period of occurrence for cynipids within galls has negative effects on the host–parasitoid relationships at higher temperature levels, thereby increasing phenological asynchrony for some parasitoids species. Reducing the development time of pupae and adults decreases the likelihood of success for some parasitoid species at higher temperature levels. We also record the extension of the gall wasp development time (approximately 15 days) at higher altitudes (linked to a lower mean temperature of approximately 1.5 °C). These results highlight how parasitization on the new hosts is dependent on the host phenology and, in the present study, is limited by the short duration of the presence of the host in galls, which could explain the considerable differences in cynipid gall wasp parasitization recorded at different altimeters.     

A nectar-secreting gall wasp and ant mutualism: selection and counter-selection shaping gall wasp phenology, fecundity and persistence

Abstract.

• 1 The natural history of a gall wasp including interactions with inquilines, parasites, and a mutualistic ant are examined. The stability of the system is described from the perspective of influences on gall wasp life history characteristics.
• 2 An exclusion experiment demonstrated that the nectar-secreting gall of Disholcaspis perniciosa mediates a mutualism with the tending ant, Formica obscuripes. Survivorship increased from 0% in the absence of ants to 25.3% in their presence, largely due to the exclusion of inquilines.
• 3 Specialized parasites, Eudecatoma spp., attacked before the ant-gall interaction began, when the developing gall was still beneath the host plant (Quercus gambellii) epidermis and ants were not in attendance. They may select for later developing gall wasps, which benefit by having fewer individuals parasitized. However, counter-selection for earlier development may result from decreased gall wasp size, decreased fecundity, and an increase in gall failures resulting from late development.
• 4 Local persistence of the gall wasp population despite increased pressure from inquilines and parasites was attributed to gall wasp escape in time due to polymorphic emergence resulting from diapause. Most individuals emerge at the end of the summer, but approximately 15% remain in the galls as prepupae for 1–5 years.

     

A plant pathogen causes extensive mortality in an invasive insect herbivore

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The dual origin of the nurse chamber in the ovarioles of the gall midge,Heteropeza pygmaea

Summary By following the lineage of the primordial germ cell and the mesoderm cells during the ontogeny of the gall midge,Heteropeza pygmaea, it has been determined that the nurse chamber of the polytrophic ovarioles of this insect is derived from the descendants of these cells. Such a dual origin for the nurse chamber of an insect ovary is the first of its kind to be reported.     

Non-random distribution among a guild of parasitoids: implications for community structure and host survival

Abstract 1. Immature stages of the gall midge, Asphondylia borrichiae, are attacked by four species of parasitoids, which vary in size and relative abundance within patches of the gall midge’s primary host plant, sea oxeye daisy (Borrichia frutescens). 2. In the current study, a bagging experiment found that the smallest wasp, Galeopsomyia haemon, was most abundant in galls exposed to natural enemies early in the experiment, when gall diameter is smallest, while the wasp with the longest ovipositor, Torymus umbilicatus, dominated the parasitoid community in galls that were not exposed until the 5th and 6th weeks when gall diameter is maximal. 3. Moreover, the mean number of parasitoids captured using large artificial galls were 70% and 150% higher compared with medium and small galls respectively, while stem height of artificial galls significantly affected parasitoid distribution. Galls that were level with the top of the sea oxeye canopy captured 60% more parasitoids compared with those below the canopy and 50% more than galls higher than the plant canopy. 4. These non‐random patterns were driven primarily by the differential distribution of the largest parasitoid, T. umbilicatus, which was found significantly more often than expected on large galls and the smallest parasitoid of the guild, G. haemon, which tended to be more common on stems level with the top of the plant canopy. 5. Large Asphondylia galls, especially those located near the top of the Borrichia canopy, were more likely to be discovered by searching parasitoids. Results using artificial galls were consistent with rates of parasitism of Asphondylia galls in native patches of sea oxeye daisy. Gall diameter was 19% greater and the rate of parasitism was reduced by almost 50% on short stems; as a result, gall abundance was 24% higher on short stems compared with ones located near the top of the plant canopy. 6. These results suggest that parasitoid community composition within galls is regulated by both interspecific differences in ovipositor length and preferences for specific gall size and/or stem length classes.     

Oviposition by introduced Ophelimus eucalypti (Hymenoptera: Eulophidae) and morphogenesis of female-induced galls on Eucalyptus saligna(Myrtaceae) in New Zealand

An Australian gall-inducing eulophid, Ophelimus eucalypti (Gahan) was first recorded on the foliage of Eucalyptus botryoides after it invaded New Zealand in 1987. It has spread throughout the eucalypt plantations in the North Island and in the northern parts of the South Island affecting several species of Eucalyptus in the section Transversaria (subgenus Symphyomyrtus). Because gall-inducing insects usually have extremely narrow host ranges, O. eucalypti that induces galls on E. saligna and E. botryoides is currently recognized as a biotype, O. eucalypti(Transversaria). Heavily galled leaves abscise from the plant. Repeated defoliation led to widespread die-back of susceptible eucalypt species in the 1990s. Female larvae of O. eucalypti induce circular, protruding galls on the leaves of E. botryoides and E. saligna, whereas the males induce pit galls on the same species. The biology of O. eucalypti females and the development of their galls are described. Adult female O. eucalypti antennate the leaf surface before inserting the ovipositor (otherwise concealed within the metasomal apex) into the young host leaf. The egg is inserted at approximately 45 degrees and discharged between differentiating palisade cells. Callus-type cells surround the egg chamber, but cytologically specialized nutritive cells appear once the egg hatches and the larva begins to feed. The gall also differentiates a multi-layered sclerenchymatous tissue around the nutritive tissue. After feeding for many months, the larva pupates and the active nutritive tissue degenerates. The adult wasp emerges after cutting an exit hole through to the outside of the gall. Abscission of heavily galled leaves results in widespread defoliation and loss of growth and vigour in susceptible trees in New Zealand.     

Diversification in the use of resources by Idarnes species: bypassing functional constraints in the fig–fig wasp interaction

Mutualisms such as the fig–fig wasp mutualism are generally exploited by parasites. We demonstrate that amongst nonpollinating fig wasps (NPFWs) parasitic on Ficus citrifolia, a species of Idarnes galls flowers and another species feeds on galls induced by other wasps killing their larvae. The galling wasp inserts its ovipositor through the fig wall into the fig cavity. The ovipositor then follows a sinuous path and is introduced through the stigma and style of the flower. The egg is deposited between the integument and nucellus, in the exact location where the pollinating mutualistic wasp would have laid its egg. Gall induction is a complex process. In contrast, the path followed by the ovipositor of the other species is straightforward: attacking a larva within a developed gall poses different constraints. Shifts in feeding regime have occurred repeatedly in NPFWs. Monitoring traits associated with such repeated evolutionary shifts may help understand underlying functional constraints. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 114–122.     

Manipulation of host plant development by the gall-midge Rhabdophaga strobiloides

Abstract. 1. The gall-midge Rhabdophaga strobiloides (O.S.) forms a gall in the apical bud of actively growing willow twigs.
2. Galls were not randomly distributed among twigs. Twigs that arose towards the distal end of the branch were much more likely to be galled. Distally located twigs also grew to greater girth than more proximally located twigs.
3. Comparisons of galled twigs with normal twigs in similar locations along their branch showed that the gall causes even greater growth in twig girth than when no gall is present.
4. The hypothesis that galled twigs draw photosynthate produced elsewhere in the plant was tested in an experiment that measured the growth of galled and normal twigs. with their leaves intact, to galled and normal twigs that were manually defoliated. Defoliation caused reduced growth in normal twigs, but galled twigs grew equally well with or without their leaves. Leaf removal had no effect on gall growth.
5. Twig diameter was positively correlated with gall diameter. Call diameter was positively correlated with larval biomass.
6. Patterns of twig and gall growth suggest that the gall-midge manipulates host plant growth and development to provide resources for growth and survival. Manipulation of the host may be an important phenomenon in the evolution of parasitic organisms.