Pollenkitt is lacking inGnetatae:Ephedra andWelwitschia; further proof for its restriction to the angiosperms

Three members of theGnetatae (Ephedra campylopoda, E. americana, Welwitschia mirabilis) were investigated by TEM and SEM with respect to their anther tapetum and pollen development. In all three species pollenkitt is lacking. The pretended pollen stickiness thus does not depend on pollenkitt. Considering former observations one can now clearly state that pollenkitt is missing in all recent gymnosperm classes (both anemophilous and ± entomophilous). Pollenkitt thus is restricted and ± omnipresent within the angiosperms, where it represents one of the most important components of the entomophily syndrome. This can be regarded as important proof for the hypothesis that the angiosperms are a single coherent phylogenetic group.     

Entwicklungsgeschichte und Ultrastruktur von Pollenkitt und Exine bei nahe verwandten entomophilen und anemophilen Angiospermensippen:Ranunculaceae,Hamamelidaceae, Platanaceae undFagaceae

Transfer of fresh pollen either by wind or by insects is strongly related to the degree of its stickiness. The sticky substances involved should be understood as pollenkitt. Ultrastructural investigations indicate that the sticky pollen of entomophilous angiosperms (in this publicationHamamelis vernalis andH. virginiana, Corylopsis platypetala) results from the deposition of usually electron-dense, homogeneous pollenkitt mainly on the outside layer of the exine. In contrast, the pollen of anemophilous angiosperms (in this publicationParrotia persica, Platanus orientalis, Thalictrum flavum andQuercus robur), but also of some entomophilous angiosperms with special pollination ecology (e.g.Aquilegia vulgaris) is rather dry and powdery. In these cases the pollenkitt often is electron-transparent and inhomogeneous and is mainly removed into the cavities of the exine. Ultrastructure and deposition of pollen and pollenkitt in ambophilous angiosperms (in this publicationThalictrum minus, Castanea sativa) are ± intermediate.

     

Entwicklungsgeschichte und Ultrastruktur von Pollenkitt und Exine bei nahe verwandten entomophilen und anemophilen Angiospermensippen derAlismataceae,Liliaceae, Juncaceae,Cyperaceae, Poaceae undAraceae

Some closely related members of the monocotyledonous familiesAlismataceae, Liliaceae, Juncaceae, Cyperaceae, Poaceae andAraceae with variable modes of pollination (insect- and wind-pollination) were studied in relation to the ultrastructure of pollenkitt and exine (amount, consistency and distribution of pollenkitt on the surface of pollen grains). The character syndromes of pollen cementing in entomophilous, anemophilous and intermediate (ambophilous or amphiphilous) monocotyledons are the same in principal as in dicotyledons. Comparing present with former results one can summarize: 1) The pollenkitt is always produced in the same manner by the anther tapetum in all angiosperm sub-classes. 2) The variable stickiness of entomophilous and anemophilous pollen always depends on the particular distribution and consistency of the pollenkitt, but not its amount on the pollen surface. 3) The mostly dry and powdery pollen of anemophilous plants always contains a variable amount of inactive pollenkitt in its exine cavities. 4) A step-by step change of the pollen cementing syndrome can be observed from entomophily towards anemophily. 5) From the omnipresence of pollenkitt in all wind-pollinated angiosperms studied one can conclude that the ancestors of anemophilous angiosperms probably have been zoophilous (i.e. entomophilous) throughout.

     

Ultrastruktur und Entwicklungsgeschichte des Pollenkitts vonEuphorbia cyparissias,E. palustris undMercurialis perennis (Euphorbiaceae)

Development, fine structure and distribution of pollenkitt is investigated inEuphorbia cyparissias, E. palustris, andMercurialis perennis. The predominantly anemophilousM. perennis produces a great amount of strictly homogeneous pollenkitt, which is deposited in the exine caves. In contrast to this and to all other angiosperms so far investigated, bothEuphorbia species produce large quantities of an extremely inhomogeneous and particular pollenkitt. Its ultrastructure is quite different, both during its development and after its deposition on the exine surface: Lipid particles with different electron density and size are wrapped in a strictly homogeneous electron transparent matrix. This can be considered as new and additional proof for the secondary entomophily ofEuphorbia.

     

Ultrastruktur und Verteilung des Pollenkitts in der insekten- und windblütigen GattungAcer (Aceraceae)

Several insect- and wind-pollinated species of the genusAcer have been investigated and compared in regard to pollen stickiness. The considerable amount of very inhomogeneous pollenkitt inA. negundo remains on the loculus wall or is deposited inside the exine cavities; thus the pollen is powdery. The small amount of non-homogeneous, granular pollenkitt inA. campestre mostly disappears into the exine cavities; only small droplets appear on the tectum surface; the pollen stickiness therefore is only moderate. On the other hand,A. pseudoplatanus andA. opalus produce an average amount of granular pollenkitt, which is deposited partially inside the exine and partially as a slender film on the tectum surface; in both the pollen is sticky.A. platanoides contains a great deal of granular and homogeneous pollenkitt; it does not only fill up the exine cavities but also extends as a thick, ± homogeneous, non-granular layer of pollenkitt over the tectum surface; therefore, the pollen is very sticky.—The characteristics of pollen agglutination together with other aspects of floral biology illustrate the wide spectrum between unequivocal entomophily and anemophily within the genusAcer: WhileA. negundo is anemophilous andA. platanoides is entomophilous, the remaining species investigated have a pollination syndrome with entomophilous and anemophilous features and are thus amphiphilous. The evolution withinAcer is tending not only towards dioecy, but also towards anemophily.

Herrn Prof. Dr.L. Geitler zum 80. Geburtstag gewidmet.     

Zur Frage der Anheftung des Pollens an blütenbesuchende Insekten mittels Pollenkitt und Viscinfäden

The attachment of pollen grains among themselves, on the loculus wall, and on flower-visiting insects is quite different in entomophilous angiosperms using pollenkitt and those using viscin threads as pollen adhesives. The sticky and viscous pollenkitt makes the pollen grains adhere, while the thin, non-elastic, non-sticky, and flexible viscin fibers fasten them like ropes on insect hairs or bristles. Nectar vomited by honey-bees, sticky stigma secretions or other additional sticky substances further improve the pollen adherence to flower-visiting insects.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Entwicklungsgeschichte und Ultrastruktur des Pollenkitts beiTilia (Tiliaceae)

The pollenkitt ofTilia platyphyllos andT. tomentosa is produced during the tetrade-stage by the tapetum plastides. After tapetum degeneration pollenkitt and accompanying substances are deposited predominantly in the intrabacular spaces of the exine, and only in small quantities on the surface of the exine itself.

     

Entwicklungsgeschichte und Ultrastruktur von Pollenkitt und Exine bei nahe verwandten entomophilen und anemophilen Sippen derOleaceae,Scrophulariaceae, Plantaginaceae undAsteraceae

Different members of the Angiosperm familiesOleaceae, Scrophulariaceae, Plantaginaceae andAsteraceae with variable modes of pollination were studied in relation to ultrastructure of pollenkitt and exine. In entomophilous representatives the mostly electron-dense and homogeneous pollenkitt is located on the outside layer of the exine, making the pollen sticky. In related anemophilous taxa the pollenkitt mostly is electrontransparent and inhomogeneous; it becomes inactive by remaining in the loculus or sinking to the bottom of the exine caves; this makes the pollen powdery. According to such criteria,Fraxinus ornus andRhinanthus serotinus are entomophilous, whileFraxinus excelsior is anemophilous. The pollen ofLathraea squamaria eventually becomes powdery and suited for wind dispersal. The broad transition field between entomo- and anemophily is demonstrated byPlantago andArtemisia. Plantago lanceolata is anemophilous, whilePl. serpentina andPl. media are amphiphilous.Artemisia nitida andA. mutellina are entomophilous,A. absinthium, A. pedemontana andA. gabriellae are amphiphilous, but tend towards entomo- (A. absinthium) or anemophily (A. gabriellae), whileA. chamaemelifolia and especiallyA. vulgaris are anemophilous.

     

Ultrastructural observations on anther tapetum development of freeze-fixed Ledebouria socialis Roth (Hyacinthaceae)

The tapetal ultrastructure of high-pressure-frozen, freeze-substituted Ledebouria socialis Roth (Hyacinthaceae) is described from the tetrad stage up to microspore mitosis. Cytoplasmic degeneration of the tapeturn occurs after microspore mitosis. During the tetrad stage and the early free-microspore stage the tapetum cells appear to be meristematic; after callose dissolution they show an intense exocytosis of polysaccharides into the anther locule. Later, the tapetum cells are characterized by abundant endoplasmic reticulum (ER). Highly osmiophilic pollenkitt precursor substances accumulate within distinct, partly irregular shaped cytoplasmic domains (“osmiophilic bodies”), which are intimately associated with the ER. It remains to be verified whether or not these bodies are derived from the ER. Because of their preservation and staining patterns the contents of these bodies are tentatively interpreted as flavonoids, one of the main pollenkitt pigments in angiosperms. Apart from these pigment bodies, there exist four other kinds of lipophilic inclusion within the anther (cells). The general aspects of lipid preservation in freeze-substituted samples are discussed. Staining with hot alcoholic phosphotungstic acid yielded good contrast of the ER and other membranes, which are often difficult to visualize in freeze-substituted, resin-embedded samples.     

A comparison of tapetal structure and function in pteridophytes and angiosperms

This article is based on a survey of literature with particular reference to the structure and functioning of the tapetum during sporogenesis in angiosperms and pteridophytes. Some original observations onSchizaea pectinata andSelaginella japonica are also included. In the past, superficial similarities in the structure and function of pollen grains and spores have dominated the research approach, but patterns of development and the functioning of the tapetum during sporogenesis in angiosperms and pteridophytes are shown to differ quite significantly and raises questions relating to homologies. It is suggested that direct comparisons may no longer be appropriate.     

Characteristics of oviposition behaviour of the pollen beetle, Meligethes aeneus on four different host plants

Oviposition behaviour of Meligethes aeneus F. (Coleoptera: Nitidulidae) is characterised and quantified on four different plant species. Six behavioural components are identified: W—walking, WA—walking with abdomen on surface, R—resting, B—biting, AOH—placing abdomen over the bite hole and OVI—oviposition. Comparison of host acceptance behaviours on Brassica napus L., Brassica juncea (L.) Czern, Brassica nigra (L.) Koch and Sinapis alba L. showed that S. alba was accepted as a host only after a long exposure to the plant. Behaviour on the Brassica species was similar, however on B. nigra beetles spent a high proportion of time actually ovipositing. We conclude that important cues for oviposition are located both on the bud surface and inside the bud.     

Interspecific variation in the defensive responses of obligate plant-ants: experimental tests and consequences for herbivory

The aggressive behavior of ants that protect plants from herbivores in exchange for rewards such as shelter or food is thought to be an important form of biotic defense against herbivory, particularly in tropical systems. To date, however, no one has compared the defensive responses of different ant taxa associated with the same plant species, and attempted to relate these differences to longer-term efficacy of ant defense. We used experimental cues associated with herbivory—physical damage and extracts of chemical volatiles from leaf tissue—to compare the aggressive responses of two ant species obligately associated with the Amazonian myrmecophyte Tococa bullifera (Melastomataceae). We also conducted a colony removal experiment to quantify the level of resistance from herbivores provided to plants by each ant species. Our experiments demonstrate that some cues eliciting a strong response from one ant species elicited no response by the other. For cues that do elicit responses, the magnitude of these responses can vary interspecifically. These patterns were consistent with the level of resistance provided from herbivores to plants. The colony removal experiment showed that both ant species defend plants from herbivores: however, herbivory was higher on plants colonized by the less aggressive ant species. Our results add to the growing body of literature indicating defensive ant responses are stimulated by cues associated with herbivory. However, they also suggest the local and regional variation in the composition of potential partner taxa could influence the ecology and evolution of defensive mutualisms in ways that have previously remained unexplored.     

Molecular evolution of the maize sex-determining gene TASSELSEED2 in Bouteloua (Poaceae)

The majority of angiosperms produce hermaphrodite flowers, while a lesser number (20–30%) produce unisexual flowers. Little is known about the molecular biology of sex-determination in angiosperms, however, a few sex-determining genes have been cloned from the model system Zea mays. One of these genes is Tasselseed2 (Ts2) which has been shown to be involved in the arrest of developing pistils in male flowers. In this study, we sequenced a putative homologue of Ts2 in species of Bouteloua, a genus in the grass subfamily Chloridoideae. We found significant genetic variation at Ts2 in Bouteloua relative to other developmental genes characterized in maize and other grass species. We also found that in Bouteluoua, Ts2 is evolving non-neutrally in the hermaphrodite-flowered Bouteloua hirsuta while no difference from neutral expectation was detected at Ts2 in the monoecious/dioecious Bouteloua dimorpha. The putatively neutral gene Alcohol Dehydrogenase1 (Adh1) was also examined for the same species of Bouteloua, and no departure from neutral expectation was detected. Our results suggest that purifying selection may be acting on Ts2 in the hermaphrodite-flowered B. hirsuta while no evidence of selection was detected at Ts2 in the monoecious/dioecious B. dimorpha.     

Self-isolation and insect pollination in the primitive angiosperms: New evaluations of older hypotheses

A review of the literature, compiled over the past 40 years suggests new directions for theories discussing the evolution of reproductive isolation and entomophily in angiosperms. Data on breeding systems suggests that the protoangiosperms may have developed gametophytic self-incompatibility. Protogyny is probably ancestral to protandry and herkogamy. Although the proto-flowers were bisexual wide variation in the number of sexual organs within proto-flowers probably led to labile sexuality and early trends towards dicliny. Beetle-pollination in extant relicts appears too specialized to represent an ancestral condition. Rather, the proto-flowers may have been generalist entomophiles incorporating some beetles,Plecoptera, thrips, micropterigid moths and proto-dipterans into their fluctuating spectra of opportunistic pollinators. Bee-pollination is probably polyphyletic in origin evolving repeatedly from angiosperms showing these generalist syndromes. There is still no correlation between primitive bees (especiallyColletidae) and the relictual angiosperms. Pollen that is usually retained within the anthers following dehiscence and the presence of staminal filaments and styles characterizes most of the flowers of those relictual angiosperms pollinated by specialized, modernApoidea.     

The chemical control of salvinia molesta (Mitchell) and some related toxicological studies

Laboratory trials were performed to screen the potentials of four herbicides — PARAQUAT, DIQUAT, TOK E-25 and DALAPON — for the chemical control of Salvinia molesta. The herbicides were applied as aerial sprays, and their phytotoxic effects were assessed by visual estimation. Paraquat and Diquat have been found to be the more promising.The toxicities of the four herbicides were tested on three aquatic organisms associated with the plant in the aquatic environment —larva of Nymphula responsalis, Rasbora trilineata and Poecilia reticulata. An attempt was made to correlate the toxicological results with the application rates of the herbicides in terms of safety factors. TOK E-25 has been found to be most toxic, the others being relatively safe.     

Physiological activities and the active constituents of potentially medicinal plants used by wild chimpanzees of the Mahale mountains,Tanzania

Potential medicinal plants for wild chimpanzees have been studied in order to discover their physiologically active compounds. Tests of the physiological activity of 3 plant species—Vernonia amygdalina, Aspilia mossambicensis, andFicus exasperata—indicate that they contain a variety of active compounds. From one species,V. amygdalina, an antitumor agent and 2 possible antitumor promoters are identified. Furthermore, steroid glucosides were isolated as the bitter substances. These structurally new compounds are expected to exhibit a number of significant physiological activities. The chemical investigation of possible medicinal plants used by chimpanzees should be helpful in recovering naturally occurring compounds of medicinal significance for human use.     

Characterization of ψM1, a virulent phage of Methanobacterium thermoautotrophicum Marburg

We have studied the biogenesis and enzymic composition of microbodies in different yeasts during adaptation of cells to a new growth environment. After a shift of cells of Candida boidinii and Hansenula polymorpha from glucose to methanol/methylamine-containing media, newly synthesized alcohol oxidase and amine oxidase are imported in one and the same organelle together with catalase; as a consequence the cells contain one class of morphologically and enzymatically identical microbodies. Similar results were obtained when Candida utilis cells were transferred from glucose to ethanol/ethylamine-containing media upon which all cells formed microbodies containing amine oxidase and catalase.However, when methanol-limited cells of H. polymorpha were transferred from media containing ammonium sulphate to those with methylamine as the nitrogen source, newly synthesized amine oxidase was incorporated only in part of the microbodies present in these cells. This uptake was confined to the few smaller organelles generally present at the perimeter of the cells, which were considered not fully developed (immature) as judged by their size. Essentially similar results were obtained when stationary phase cells of C. boidinii or C. utilis — grown on methanol and ethanol plus ammonium sulphate, respectively — were shifted to media containing (m)ethylamine as the nitrogen source. These results indicate that mature microbodies may exist in yeasts which no longer are involved in the uptake of matrix proteins. Therefore, these yeasts may display heterogeneities in their microbody population.     

Invasive Grass Alters Litter Decomposition by Influencing Macrodetritivores

The results of nitrogen (N) fertilization experiments have shown inconsistent rates of plant litter decomposition, a phenomenon that may be explained by dispropotionate influence of animal detritivores (macro-detritivores) on litter mass loss versus that of microbial decomposers, whose activity may be dependent on inorganic N. In turn, macrodetritivores may be influenced by plant species composition via their selection of optimal food resources and habitats. In our experiment, fertilizer had no apparent effect on litter decomposition, suggesting that microbial decomposers did not use the additional inorganic N and/or that macrodetritivores had a greater influence on decomposition. Manipulation of macrodetritivores suggested that plant species composition—dominated in this study by Festuca arundinacea, an exotic, invasive grass, and Aster ericoides, a native forb—caused shifts in detrivore communities and/or feeding patterns that tended to increase litter mass loss. Canopy cover of F. arundinacea and A. ericoides ranged from 0% to 11%, suggesting that low-intensity invasion may produce significant changes in ecosystem function, such as decomposition.