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So me species of the paleotropical tree genus Macaranga (Euphorbiaceae) live in elose association with ants. Thc genus comprises the full range of species from those not regularly inhabited by ants to obligate myrmecophytes. In Malaysia (peninsular and Borneo) 23 ofthe 52 species areknown to be ant-associated (44%). The simplest structural adaptation of plants to attract ants are extrafloral nectaries. We studied the distribution of extraflural nectaries in the genus Macaranga to assess the significance of this character as a possible predisposition for the evolution of obligate myrmecophytism. All species have marginal glands on the leaves. However, only the glands of nonmyrmecophytic species function as nectaries, whereas liquids secreted by these glands in myrmecophytic species did not contain sugar. Some non-myrmecophytic Macaranga and transitional Macaranga species in addition have extrafloral nectaries on the leaf blade near the petiole insertion. All obligatorily myrmecophytic Macaranga species, however, lack additional glands on the lamina. The non-myrmecophytic species are visited by a variety of different ant species, whereas myrmecophytic Macaranga are associated only with one specific ant-partner. Since these ants keep scale insects in the hollow sterns, reduction of nectary production in ant-inhabited Macaranga seems to be biologically significant. We interpret this as a means of (a) saving the assimilates and (b) stabilization of maintenance of the association's specificity. Competition with other ant species for food rewards is avoided and thereby danger ofweakening the protective function ofthe obligate antpartner for the plant is reduced. A comparison with other euphorb species living in the same habitats as Macaranga showed that in genera in which extrafloral nectaries are widespread, no myrmecophytes have evolved. Possession of extrafloral nectaries does not appear to be essential for the development of symbiotic ant-plant interactions. Other predispositions such as nesting space might have played a more important role.
Zanthoxylum myriacanthum, a small Rutaceous tree growing mainly in secondary hill forests in SE Asia, is a true myrmecophyte. It possesses stem domatia in the form of hollow branches with slitlike openings. Branch hollows and entrance slits are produced by the plant itself through pith degene~.tion ?u.d growth proceSses. If the entrance is not kept open by ants it closes again by growth ol the surrounding tissue after some time. The domatia are colonized opportunistic ally by different arboreous ants, e.g. Crematogaster and Campono tus. Additionally many small extrafloral nectaries are found on the leaflets of Zanthoxylum myriacanthum. Judging from herbarium studies and literature records at least four more true ant trees are found in the genus Zanthoxylum namely Z. rhetsa in SE Asia, Z. conspersipunctatum, Z. pluviatile and Z. vinkii in New Guinea. We could not confirm ant inhabitation in Drypetes pendula (Euphorbiaceae) on the Malay Peninsula, which has also been recorded to be an anttree.
The production of extrafloral nectar and food bodies plays an important role in many tropical ant-plant mutualisms. In Malaysia, a close association exists between ants and some species of the pioneer tree genus Macaranga (Euphorbiaccac). Macaranga is a very diverse genus which exhibits all stages ofintcraction with ants, from facultative to obligatory associations. The ants nest inside the hollow inlcrnodes and reed mainly on food budies provided by the plants. Food body production had previously been reported only in myrrnecophytic Macaranga species, where it is usually coneentrated on protected parts or the plants such as recurved stipules. We found that non-myrmecophytic Macaranga species also produce food bodies on leaves and stems, where they are collected by a variety or ants. Levels of food body production differ between facultatively and obligatorily ant-associated species but also among the various non-myrmecophytes. This may he rdated to the degree of interaction with ants. Food body production starts at a younger age in the myrmccophytic species than in the transitional or non-myrmcccophytic Macaranga. Although food bodies of the non-inhabited Macaranga species are collected by a variety of ants, there is nu evidence of association with specific ant species. Our observations suggest that food bodies enhance the evolution of ant-plant interactions. Production of food bodies alone, however, does not appear to be the most important factor for the development of obligate myrmccopllytism in Macaranga.
Lampbrush chromosomes of amphibian oocytes were isolated in the presence of near-physiological salt concentrations, to preserve their native state, and studied by electron microscopy of ultrathin s~dions. The transcriptional state of the lampbrush chromosomes was experimentally modulated by incubating the oocytes for various time periods in medium containing actinomycin D. The observations show that the structure of the lateral loops changes rapidly in response to alterations in transcriptional activity. During decreasing transcriptional activity and reduced packing density of transcripts, the chromatin axis first condensed into nucleosomes and then into an approximately 30 nm thick higher order chromatin fiber. Packaging of the loop axis into supranucleosomal structures may contribute to the foreshortening and retraction of the loops observed during inhibition of transcription and in later stages of meiotic prophase. The increasing packing density of the DNA during the retraction process of the loops could also be visualized by immunofluorescence microscopy using antibodies to DNA. The dependence of the loop chromatin structure on transcriptional activity is discussed in relation to current views of mechanisms involved in gene activation.
The distribution of DNA in Ehrlich tumour cell nucleoli was investigated by means of an immunocytochemical approach , involving a monoclonal antibody directed against double- and single-stranded DNA. Immunolabelling was performed . either before or after the embedding process. The postembedding labelling method allows better ultrastructural preservation than the preembedding labelling method. In particular, the various nucleolar components are well preserved and identifiable. In the nucleolus, labelling is particularly concentrated over the perinucleolar chromatin and over its intranucleolar invaginations, which penetrate the nucleolar body and often terminate at the fibrillar centres. In addition, aggregates of gold particles are found in the fibrillar centres, preferentially towards the peripheral regions. By contrast, the dense fibrillar component is completely devoid of labelling. The results seem to indicate that DNA containing the rDNA genes is located in the fibrillar centres, with a preference for the peripheral regions. This finding suggests that transcription of the rDNA genes should occur within the confines of the fibrillar centre, probably close to the boundary region of the surrounding dense fibrillar component. The results are discussed in the light of present knowledge of the functional organization of the nucleolus.
Small nucleolus-related bodies which occur in the nUcleoplasm of " micronuclei" lacking nucleolar organizers have been studied by immunofluorescence microscopy. These bodies stained specifically with three different antibodies directed against proteins that are normally associated with the dense fibrillar component of functional nucleoli, but not with antibodies specific for certain proteins of the granular component or the fibrillar centers. Our data show that, in the absence of rRNA genes, the various constituent proteins characteristic of the dense fibrillar component spontaneously assemble into spherical entities but that the subsequent fusion of these bodies into larger structures is prevented in these micronuclei. The similarity between these nucleolus-related bodies of micronuclei and the prenucleolar bodies characteristic of early stages of nucleologenesis during mitotic telophase is discussed.
The present article had originally been conceived as a review on endomembranes, the plasma membrane, and the major product of membrane-bound activities, the cell wall material. However, limitations of space and the cascading number of pertinent literature articles made it necessary to confine this to one group of membranes and one type of cell wall components. Therefore, we shall begin our survey on the biochemical and cytological aspects of membranes by a review of the class of the pore complex bearing endomembranes, i.e. the nuclear envelope and the annulate lamellae (AL). Next year the membranes of the endoplasmic reticulum and the dictyosomes will be dealt with in conjunction with a discussion of the various intracellular vesicles, the tonoplast and the plasmalemma.
In order to investigate the chemical composition of the nuclear pore complexes isolated nuclei from mature Xenopus laevis oocytes were manually fractioned into nucleo· plasmic aggregates and the nuclear envelopes. The whole isolation procedure takes no more than 60- 90 sec, and the pore complexes of the isolated envelopes are well preserved as demonstrated by electron microscopy. Minor nucleoplasmic and cytoplasmic contaminations associated with the isolated nuclear envelopes were determined with electron microscopic morphometry and were found to be quantitatively negligible as far as their mass and nucleic acid content is concerned. The RNA content of the fractions was determined by direct phosphorus analysis after differential alkaline hydrolysis. Approximately 9% of the total nuclear RNA of the mature Xenopus egg was found to be attached to the nuclear envelope. The nonmembranous elements of one pore complex contain 0.41 X 10- 16 g RNA. This value agrees well with the content estimated from morphometric data. The RNA package density in the pore complexes (270 X 10- 15 g/fJ-3) is compared with the nucleolar, nucleoplasmic and cytoplasmic RNA concentration and is discussed in context with the importance of the pore complexes for the nucleo-cytoplasmic transport of RNA-containing macromolecules. Additionally, the results of the chemical analyses as well as of the 3H-actinomycin D autoradiography and of the nucleoprotein staining method of Bernhard (1969) speak against the occurence of considerable amounts of DNA in the nuclear pore complex structures.
Electron opaque cross-bridge structures span the inter- and intracisternal spaces and provide membrane-to-membrane connections between adjacent cisternae of dictyosomes of pollen tubes of Clivia and Lilium. Additionally, the classic intercisternal rods, characteristic of intercisternal regions near the maturing face of dictyosomes, are connected with the adjacent membranes through similar cross-bridge elements. We suggest that these structural links are responsible for maintaining the flattened appearance of the central parts of Golgi apparatus cisternae as well as for the coherence of cisternae within the stack. Observations on other plant (e.g. microsporocytes of Canna) and animal cells (e.g. rodent liver and hepatoma cells, newt spermatocytes) show that such an array of membrane cross-links is a universal feature of Golgi apparatus architecture. The cross-bridges appear as part of the complex "zone of exclusion" which surrounds dictyosomes, entire Golgi apparatus and Golgi apparatus equivalents in a variety of cell types.
The disintegration of the nuclear envelope has been examined in nuclei and nuclear envelopes isolated from amphibian oocytes and rat liver tissue, using different electron microscope techniques (ultrathin sections and negatively or positively stained spread preparations). Various treatments were studied, including disruption by surface tension forces, very low salt concentrations, and non ionic detergents such as Triton X-lOO and Nonidet P-40. The high local stability of the cylinders of nonmembranous pore complex material is emphasized. As progressive disintegration occurred in the membrane regions, a network of fibrils became apparent which interconnects the pore complexes and is distinguished from the pore complexassociated intranuclear fibrils. This network might correspond to an indistinct lamella, about 15 - 20 nm thick, located at the level of the inner nuclear membrane, which is recognized in thin sections to bridge the interpore distances. With all disintegration treatments a somewhat higher susceptibility of the outer nuclear membrane is notable, but a selective removal does not take place. Final stages of disintegration are generally characterized by the absence of identifiable, membrane- like structures. Analysis of detergent-treated nuclei and nuclear membrane fractions shows almost complete absence of lipid components but retention of significant amount of glycoproteins with a typical endomembrane-type carbohydrate pattern. Various alternative interpretations of these observations are discussed. From the present observations and those of Aaronson and Blobel (1,2), we favor the notion that threadlike intrinsic membrane components are stabilized by their attachment to the pore complexes, and perhaps also to peripheral nuclear structures, and constitute a detergent-resistant, interpore skeleton meshwork.