ERUMPENT CEPHALODIA,AN APPARENT CASE OF PHYCOBIAL INFLUENCE ON LICHEN MORPHOLOGY1,2,3

On the upper surface of some specimens of the lichen Lobaria cf. crosa (Eschw.) Nyl. there are shrub-like growths which have a morphology similar to that of a free-living fruticose lichen, Polychidium umhausense (Auersw.) Henss., and contain a cyanophyte phycobiont, Nostoc. A few growths also contain scattered colonies of a chlorophyte phycobiont, in which case the lichen tissue locally assumes the foliose form characteristic of the parent Lobaria thallus. The differentiation of dorsiventral lichen cortices and the formation of a lax medulla and distinct algal layer are correlated with the presence of the green phycobiont. The lichen substances atranorin, gyrophoric acid, and 4-0-methylgyrophoric acid occur in both, the foliose L. erosa thallus and the fruticose tissue. It is suggested that the fruticose structures are erumpent cephalodia which are derived from the outgrowth of internal, cephalodia and should not be considered, to be epiphytic colonies of the lichen Polychidium.     

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.     

ETIOLOGY AND DEVELOPMENT OF HYPERPLASIA INDUCED BY STREBLONEMA SP. (PHAEOPHYTA) ON MEMBERS OF THE LAMINARIALES (PHAEOPHYTA)1

Filaments of Streblonema sp. isolated from hyperplasia (galls) on Nereocystis luetkeana (Mert.) Post. et Rupr. induced similar gall growths when inoculated on young sporophytes of Nereocystis luetkeana, Macrocystis integrifolia Bory, and Laminaria japonica Aresch. Filaments were irregularly branched and uniseriate with cells 5–8 μm in diameter and 10–30 μm long. Growth under varied cultured conditions produced unilocular sporangia. Zoospores released from the sporangia germinated into identical filaments which also formed unilocular sporangia. Gall tissue originated from the innermost cells of the epidermal meristematic zone. When infected these cells divided in an irregular and unorganized manner. The resulting structure was a pronounced departure from normal morphology.     

Reiterative production and deformation of cell walls in expanding thallus nets of the lichen ramalina menziesii (Lecanorales,Ascomycetes)

The thallus of the lichen Ramalina menziesii Tayl. is composed of net-like units that develop by diffuse expansion of perforated tissue produced at the net apex. Study of net tissue with transmission electron microscopy reveals that the cortical cells are surrounded by a succession of cell walls alternating with layers of an electron-transparent matrix substance. In the course of thallus growth the cortical cell walls are continually deformed and new ones constructed. The deposition of new walls and matrix layers displaces the older walls centrifugally from the cell. Electron-dense boundaries develop at the interfaces among cells where the remains of the oldest walls are compressed against those of neighboring cells. As new branch cells are inserted through the concentric accumulations, the dense boundaries appear to enclose fascicles of cells, visible in cross section with light microscopy. Cortical organization in Ramalina menziesii is contrasted with that reported in other lichens, and a functional relationship to diffuse growth of the thallus is suggested.     

APICAL DOMINANCE,POLARITY, AND ADVENTITIOUS GROWTH IN MARCHANTIA POLYMORPHA

Benzylaminopurine, indoleacetic acid, and the auxin inhibitors transcinnamic acid, triiodobenzoic acid, and dinitrophenol were employed to elucidate the role of apical dominance and hormones during regeneration of thalli and gemmae of Marchantia polymorpha L. The cytokinin suppressed normal gemma germination and led to the development of nodular, callus-like growths. When removed from the influence of benzylaminopurine, the site and magnitude of normal thallus outgrowths varied with the length of time that the tissue had remained on the cytokinin-containing medium. This aberrant germination was not influenced by the incorporation of indoleacetic acid into the medium. Exogenous auxin neither accelerated nor inhibited the regeneration of normal thallus growth on excised vegetative discs. Transcinnamic acid and dinitrophenol inhibited regeneration. Auxin reversed this suppression. Triiodobenzoic acid did not significantly affect regeneration. Autoradiographs demonstrated a pronounced accumulation of labelled auxin in the midribs and the acropetal regions of excised thallus discs. This evidence suggests that there is an endogenous, basipetal auxin gradient in Marchantia; that the maintenance of this gradient is vital to normal growth and regeneration of the thallus; and that high endogenous concentrations of cytokinin destroy this polarity by increasing the auxin-synthesizing capacity of the tissue.     

INDOLE ACETIC ACID OXIDASE AND PEROXIDASE ACTIVITIES IN NORMAL AND CROWN GALL TISSUE CULTURES OF PARTHENOCISSUS TRICUSPIDATA

Lipetz , Jacques , and Arthur W. Galston . (Yale U., New Haven.) Indole acetic acid oxidase and peroxidase activities in normal and crown gall tissue cultures of Parthenocissus tricuspidata. Amer. Jour. Bot. 46(3) : 193-196. Illus. 1959.—Normal and crown gall cells of P. tricuspidata grown in pure culture were examined for IAA oxidase and peroxidase activities. No IAA oxidase activity could be demonstrated in dialyzed or undialyzed homogenates of either tissue; however, crown gall tissue, but not normal tissue, was found to produce an extracellular IAA oxidase which required Mn⁺⁺ and DCP as co-factors. Normal tissue, but not crown gall tissue was found to contain high levels of substances which spared IAA from destruction by a pea IAA oxidase preparation. Peroxidase activity was found to be higher in normal than in crown gall homogenates, but crown gall tissue released considerably more peroxidase into the external medium. The differences in the auxin requirements and growth rate between normal and crown gall cells appear not to be easily explicable in terms of differential auxin destruction.     

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.     

Meristoderm in Turbinaria conoides (Fucales,sargassaceae)

In Turbinaria conoides (J. Agardh) Kützing, the hapteron, thallus and receptacle meristoderm cells are columnar, polarized and have thick longitudinal walls. In the thallus apical cavity, however, these cells are elongate and possess cap-like structures and thin, longitudinal walls. The meristoderm cells, except for those present at the apical cavities and ostiole regions, are overarched by 1–5 extracellular layers that are deposited in an orderly manner.The histochemistry of the meristoderm suggests that its cell walls contain alginic acid, cellulose and sulphated polysaccharides. The extracellular layers and the materials in the cap-like structures contain a mixture of alginic acid and sulphated polysaccharides, and little cellulose. The possible role of meristoderm cells lining the apical cavity of the thallus and that of extracellular layers is discussed.     

Lithophyllum cuneatum sp. nov. (Corallinaceae, Rhodophyta), a new species of non-geniculate coralline alga semi-endophytic in Hydrolithon onkodes and Neogoniolithon sp. from Fiji, South Pacific

A new species of semi-endophytic coralline alga, Lithophyllum cuneatum (Corallinaceae: Lithophylloideae), is described from Fiji. The species is characterized by a wedge-like thallus that is partially buried in the thallus of the host coralline, Hydrolithon onkodes (Heydrich) Penrose et Woelkerling or occasionally Neogoniolithon sp., and that appears at the surface of the host as a small pustule that is usually paler in color than the host. The thallus consists of erect filaments that are derived from a single cell. The basal cell, when visible, is non-palisade, and areas of bistratose margin are absent. Cells of contiguous erect filaments are joined by secondary pit connections. Epithallial cells are present in 2–3 layers, and individual trichocytes are common. Gametangial plants are dioecious. Male conceptacles have simple spermatangial systems that are confined to the floors of their elliptical chambers. Carposporangial conceptacles contain 5–8 celled gonimoblast filaments that are borne at the margin of a more-or-less discoid fusion cell, and so occupy the periphery of the elliptical conceptacle chambers. Tetrasporangial conceptacles are uniporate, with roofs formed from peripheral filaments, and chambers lack a central columella of sterile filaments. Despite its semi-endophytic nature, haustorial cells are absent, and plastids and pigmentation are present.     

Evidence for the Presence of a Readily Transmissible Oncogenic Principle in Crown Gall Teratoma Cells of Tobacco

Teratomas result when totipotent tobacco cells conditioned by wounding are exposed to the moderately virulent T37 strain of the crown gall bacterium. Once transformation is accomplished, teratoma cells continue to proliferate autonomously in the absence of the inciting bacterium. The present report demonstrates that teratoma tissues regularly induce the production of new non-selflimiting growths when grafted onto suitable host plants. The newly induced tumours are derived from the normal cells of the host. It is concluded that teratoma cells contain an oncogenic principle in an active form. Whether this principle is identical with that elaborated by the bacteria, is not as yet known.     

THALLUS ORGANIZATION AND DEVELOPMENT IN THE FRUTICOSE LICHENASPICILIA CALIFORNICA,WITH COMPARISONS TO OTHER TAXA

Thallus organization is examined inAspicilia californicaRosentreter, a fruticose lichen known from several localities in central and southern California. The sprawling, terete thallus branches possess a dense central medulla of thick-walled, longitudinally oriented fungal cells. This central tissue emerges at branch apices to form a darkly pigmented fungal tip. Thallus development involves the apical extension of the tip to produce a fungal tissue over which a cylindrical algal layer and cortex will eventually be formed. Apical branches are initiated by furcation entirely within the fungal tip. Lateral branches, emerging from the lichenized thallus, arise as a divergent bundle of elongate fungal cells originating in the medulla. The photobiont appears to play no direct role in initiation of apical or lateral branches. It is concluded that thallus development inA. californicaoccurs with a relatively low degree of synchrony between mycobiont and photobiont growth, similar to the pattern observed in crustose lichens with prothallic growth. A rather similar type of thallus organization is observed inA. hispida, although in that species mycobiont growth and branch initiation appear to be somewhat more closely associated with algal cell proliferation. A squamuloseAspiciliafrom central Spain produces rhizomorphs that may sometimes become invested with an algal layer and cortex, resembling the thallus axes ofA. californica.     

CYANOPHYTE CEPHALODIA IN THE LICHEN GENUS NEPHROMA

The lichens, Nephroma expallidum (Nyl.) Nyl. and N. arcticum (L.) Torss., consistently have at least two symbionts in a single thallus: a green alga in the algal layer and a blue-green alga in the internal cephalodia. The cephalodia originate from algal cells in contact with the lower surface of the lichen, in the zone of rhizine formation. The rhizines surround the epiphytic algal colony and form a second cortical layer; following dissociation of the original lower cortex, further growth of the two organisms results in the cyanophyte colony being enveloped by a compact layer of fungal tissue and positioned in the lichen medulla. The colony may eventually assume a superior or inferior position in relation to the lichen thallus, depending in part on the lichen species. Nephroma anticum may have two distinct morphological forms of blue-green algae in the same thallus and occasionally in the same cephalodium. It appears that the relationship that exists between the cephalodial algae and the lichen thallus is antagonistic and results, in some cases, in the exclusion of the green algal layer and death to the cephalodial cyanophytes.     

Whitening,thallus decay and fragmentation in Gracilaria chilensis associated with an endophytic amoeba

Whitening of Gracilaria chilensis, accompanied by tissue softening and thallus fragmentation, was found to be associated with the presence of an endophytic amoeba. Although the symptoms developed originally in green mutant thalli, subsequent infections in the laboratory also affected normal, wild-type G. chilensis. Ultrastructural evidence indicates that the amoebae perforate the host cell walls of both cortical and medullary cells and digest their protoplasm. Feeding by the amoeba appears to involve both phagocytosis and enzymatic digestion of the host tissue. Destruction of the host tissue resulted in large cavities first, followed by thallus fragmentation. No other organism was found during the early stages of thallus invasion by the amoeba, although bacteria may appear once the amoeba reaches the inner tissues of the host.     

Studies on Metamastophora (Corallinaceae,Rhodophyta). I. M. flabellata (Sonder) Setchell: Morphology and anatomy

A morphological-anatomical study of Australian populations of Metamastophora flabellata (Sonder) Setchell, the type species of Metamastophora (Corallinaceae, Rhodophyta), has revealed that the primarily erect or ascending non-geniculate thallus possesses a dorsi-ventral organization of tissues. All conceptacles are uniporate and arise dorsally. Two distinct vegetative meristems occur: an apical primary meristem from which hypothallial cells are produced basipetally and a sub-epithallial secondary meristem which generates perithallial cells basipetally and secondary epithallial cells acropetally. Primary epithallial cells arise from divisions of subapical hypothallial cells. In younger parts, tissues are produced only dorsal to the hypothallium; in veins and stipes, tissue production occurs both dorsal and ventral to the hypothallium. Mature tetrasporic conceptacles contain peripheral tetrasporangia with zonately divided contents and a central sterile columella. Gametic conceptacles produce fertile tissue across the entire conceptacle chamber floor. After fertilization, the zygotic nucleus or a derivative is transferred (presumably) to an auxiliary cell through cells of the carpogonial branch; no tubular transfer siphon develops. Mature fusion cells are composed of the amalgamated supporting cells of carpogonial branches and are initiated from a single supporting cell which functions as an auxiliary cell. Unbranched 3–4 celled gonimoblast filaments arise from the fusion cell, do not become connected to other cells, and produce terminal carposporangia. Results from this study have led to a redefinition of hypothallium and perithallium in relation to meristems rather than substrate. In addition, carposporophyte ontogeny in the Corallinaceae is considered in terms of the presumed mode of transfer of the zygotic nucleus to the fusion cell, the extent of fusion cell development, and gonimoblast filament production in relation to auxiliary cells and fusion cells.     

ANATOMY OF SOME LEAF GALLS OF ROSA WOODSII (ROSACEAE)

Infection of Rosa woodsii by some members of the order Hymenoptera results in neoplasmic outgrowths on the leaves. One type of outgrowth produces a spherical swelling (leaf gall) while the other has extensive hair-like proliferations (hairy gall). The anatomy and ultrastructure of these galls were examined by light microscopy and transmission electron microscopy. The leaf gall cells were considerably larger than normal cells, lacked well-developed chloroplasts and were loosely arranged with prominent intercellular spaces. Vascular bundles were scattered throughout the gall tissue. The upper three cell layers of the leaf gall tissue resembles a periderm, having many suberin lamellae. The suberin lamellae were often traversed by pores which may represent incomplete plasmodesmata. Phenolic compounds were commonly seen both in the normal and gall cells. A layer of internal cells of the hairy galls have remarkably thickened cell walls, presumably due to the deposition of cellulosic substances. Unlike leaf galls, the epidermal cells of the hairy galls were not heavily cuticularized and no periderm was found. The hair-like outgrowths present on the outer surface of these galls had a central vascular bundle. The epidermis of the outgrowths also had thickened cell walls, and trichomes occurred on the outer surface. The structural modifications brought about by the insect invasion in these two galls are compared and their roles in gall formation are discussed.     

Histopathology of bacterial gall of Japanese Wistaria (Wistaria floribunda)

Tumor development in wistaria gall caused byErwinia milletiae, (Kawakami and Yoshida) Magrou was observed under a microscope. Hypertrophied cells were observed near wounds 3 days after inoculation. Active cell divisions subsequently occurred in the hypertrophied cells resulting in many daughter cells with thin cell walls, large intensely stained nuclei, and less vacuolated, dense cytoplasm. Hyperplasia occurred often in the concentric arrangements around large bacterial fissures. These cell groups, which were commonly found in phloem, were considered to be the elementary unit tissue of the tumor. With the development of each unit tissue, the surrounding parenchyma tissues appeared to be markedly compressed. The hyperplastic cells near cambium differentiated into the tracheal elements which developed later into vascular bundles extending to the gall tissues. When the tumor tissues further proliferated with a rapid increase in dimensions, the tumor finally erupted out of the periderm of the stem forming a visible gall. The gall tissues were quite different from those of the root nodules caused byRhizobium sp. in that they had an absence of periderm, the full development of the tracheal elements, the diversity of cells in both size and shape, and the multiplication of the pathogenic bacterium in the intercellular spaces.     

DEVELOPMENTAL ANATOMY OF THE HORNED OAK GALL INDUCED BY CALLIRHYTIS CORNIGERA ON QUERCUS PALUSTRIS (PIN OAK)

The horned oak gall forms on twigs of Quercus palustris Muench. and is initiated when the wasp Callirhytis corrigera O.S. oviposits into the periderm or cortex of the twigs. Injury to phellogen as a consequence of oviposition results in dedifferentiation of phelloderm and underlying phloem tissue to form a wound-response phellogen. The wound-response phellogen encircles and compartmentalizes the wasp ovum and is continuous with the normal stem phellogen. This tissue arrangement forms the framework for the ensuing stages of gall development. Development of the insect larva coincides with the formation of a gall-matrix phellogen. This additional meristem is derived from locally hyperplastic stem phellogen where adjacent to the wound-response phellogen. The essentially parenchymatous, roughly spherical body of the gall, the gall matrix, is formed chiefly from derivatives of the gall-matrix and wound-response phellogens. In the immediate area of the wasp larva, the wound-response phellogen produces cells toward the larva that differentiate to form a vascularized, externally sclerified, horn-shaped chamber containing in its base the developing insect larva. The involvement of the vascular cambium is apparent during intermediate gall development stages. Vascular strands, produced transverse to the stem axis, penetrate the gall-matrix tissue and terminate near the base of the horn-like larval chamber. Vessel elements in the region of the gall that are not included in this tissue alteration often become heavily tylosed. Disruption of the vascular tissue through the galled area appears to be a primary cause of twig death.     

The anatomy of Neotenophycus ichthyosteus gen. et sp. nov. (Rhodomelaceae,Ceramiales), a bizarre red algal parasite from the central Pacific

A minute parasite of Neosiphonia poko (Hollenberg) Abbott from a shallow lagoon on the central-Pacific Johnston Atoll is described as Neotenophycus ichthyosteus Kraft et Abbott, gen. et sp. nov. The infective parasite cell first connects to a central-axial cell of the host, then emerges from between host pericentral cells at a node before dividing into a three- or four-celled primary axis. Epibasal cells of the parasite divide to form three pericentral cells whose derivatives produce a globular head on the basal cell and on which reproductive structures differentiate almost immediately. Trichoblasts on any life-history stage are completely lacking. Spermatangia are borne on mother cells across the whole thallus surface. Procarps consist of four pericentral cells that encircle a subapical fertile-axial cell in an ampullar configuration, one of the pericentral cells serving as the supporting cell and bearing a four-celled carpogonial branch and a single sterile cell. Diploidization results in a longitudinal/concave division of the auxiliary cell and formation of an arching linear series of inner gonimoblast cells, each dividing toward the thallus surface into gonimoblast filaments of very narrow, horizontally aligned cells terminated by initially monopodial, later by sympodial, carposporangia, the whole of the mature female gametophyte consisting of an amalgam of several cystocarps within a lax jacket of sterile gametophytic tissue. Tetrasporophytes are composed of lobes of pericentral-cell-derived filaments, each axial cell of which is ringed by three pericentral cells producing tetrahedral tetrasporangia enclosed by two pre-sporangial cover cells. Affinities of the new genus are discussed and comparison is made particularly to the enigmatic parasite Episporium centroceratis Möbius. It is concluded that relationships with any previously described tribe are so remote or obscure that the new tribe Neotenophyceae should be proposed for it.     

THE FINE STRUCTURE OF SIEVE ELEMENTS OF NEREOCYSTIS LÜTKEANA

The sieve elements of Nereocystis from the base of phylloids contain numerous small vesicles, cytoplasm, ribosomes, and the usual organelles and membrane systems, including nuclei, plastids, mitochondria, dictyosomes, and endoplasmic reticulum. They have a thick secondary wall layer which is deposited along the longitudinal walls and at the sieve plate excluding the sieve pores. The sieve pores range in diameter from 100 to 400 nm and are lined by plasmalemma. The sieve elements from the hollow basal parts of the pneumatocyst show essentially the same features but have larger and fewer vesicles, relatively little cytoplasm, larger sieve pores, 400–900 nm in diameter, and may lack a nucleus. In old sieve elements there are large deposits of callose on the sieve plate and along the longitudinal wall; the vesicles seem to break down, and the protoplast appears necrotic. It is concluded that the trumpet hyphae and sieve tubes are basically the same type of cell, and that the trumpet-shape of the sieve elements is due to their passive stretching during extension growth of the organ in which they occur. There are minor but significant differences among the sieve elements from different regions of the thallus which may reflect possible levels of structural specialization of the sieve elements within the same plant.     

Carnivores and carotenoids are associated with adaptive behavioural divergence in a radiation of gall midges

1. Adaptive divergence in sympatry is supposed to be inhibited by the homogenizing role of gene flow. However, studies continue to uncover examples of sympatric divergence. In this study, two divergent phenotypes in a complex of four syntopic gall midge morphotypes [nominally Asteromyia carbonifera Osten Saken, Diptera: Cecidomyiidae: Alycaulini] are characterised. The first is a behavioural phenotype governing within‐host tissue preference and the second is a trait governing accessory‐gland carotenoid quality and quantity. 2. One gall morphotype (crescents) lay most of their eggs on mature tissue whereas the other three gall morphotypes oviposit only on young emerging leaves. Ecological maintenance of this divergent trait appears to be driven by enemy‐reduced space. That is, nearly 40% of the crescent morphotype galls that develop high on the plant are attacked by the egg parasitoid Platygaster solidaginis Ashmed, whereas those low on the plant are relatively protected. 3. All morphotypes contain carotenoids in their accessory glands, but the quality and quantity of these pigments differs significantly between the morphotypes and is positively associated with the probability of parasitism by P. solidaginis. 4. Larval salivary glands also contain carotenoids and the plant hormone abscisic acid, which in plants is synthesized from carotenoid precursors and is involved in regulating plant defences. This hormone may facilitate gall development and influence gall morphology. 5. Ecological fitness trade‐offs between carotenoids, parasitoid attack, and plant resistance may be fostering adaptive divergence in ovipositional phenotypes and sympatric speciation in this complex of gall midge morphotypes.