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1.
Summary The fine structure of the various hormone-producing cell types (with the exclusion of the prolactin cells) in the pituitary gland (pars distalis) of migratory sockeye salmon is described. All fish were in an advanced stage of sexual maturation. In the proximal pars distalis five cell types were distinguished: growth hormone cells, ACTH cells, gonadotrops, vesicular cells, and chromophobe cells. Gonadotrops were also found throughout the rostral pars distalis. A conspicuous feature of the gonadotrops was the presence of two kinds of secretory inclusions: small electron-dense granules (200–375 m) and large, relatively electron-translucent globules (400–2 000 m). The large vesicular cells, so called because of their conspicuous vesicular endoplasmic reticulum, were numerous and often appeared to contain some small granules. It is argued that they may represent a second type of gonadotropic cell, which, in earlier stages of gonad development, contains many granules but becomes largely degranulated near the time of reproduction when the other gonadotrops (globular gonadotrops) abound. The chromophobes, which were smaller and far less abundant than the vesicular cells, also appeared to contain small granules (120–280 m). They are probably thyrotrops.The assistance of Mr. S. Killick, of the International Pacific Salmon Fisheries Commission, who helped in the collection of salmon, is gratefully acknowledged.  相似文献   

2.
Summary Rapid freeze-substitution fixation was employed in immunocytochemical studies on the localization of LH and FSH in the typical gonadotrophs of the anterior pituitary in the untreated male rat; a modification of a recently described ferritin antibody method (Inoue et al. 1982) was used in these studies. It was shown that rapid freeze-substitution fixation provides good preservation not only of the ultrastructure but also of the antigenicity. Both LH and FSH were clearly demonstrated in the same gonadotrophic cells, but the subcellular localization of these gonadotrophins differed: (i) LH was mainly located in small secretory granules, 250–300 nm in diameter; (ii) FSH was mainly present in large secretory granules, up to 500 nm in diameter. In the pituitary gland of the adult male rat, all gonadotrophs that react to antibodies against gonadotrophins are characterized by small and large secretory granules. Other types of cells of the anterior pituitary containing either small secretory granules or resembling corticotrophs with secretory granules assembled at cell periphery did not react to either anti-LH beta or anti-FSH beta serum.For light microscopy, the peroxidase antibody method was used. All of the gonadotrophin-positive cells contain both LH and FSH. None of the pituitary cells reacted to antibody against only one gonadotrophin. However, some cells are LH-rich while other cells are FSH-rich.  相似文献   

3.
Summary The ultrastructure of the intermediate lobe of the hypophysis was studied in Anolis carolinensis with the use of a threefold aldehyde fixative. Lizards with a brown skin were selected. The possibility of two types of secretory cells is discussed; neither cell type is innervated. Type I cells are rarely found and contain dense granules approximately 0.3 m in diameter; Type II cells vary widely in secretory activity. Most of the Type II cells contain a large number of dense secretory granules (up to about 1.3 m in diameter) almost filling the cytoplasm. Rough endoplasmic reticulum (RER), Golgi apparatus and mitochondria are poorly developed. Only some of these cells show signs suggesting a high secretory activity, namely a well developed RER, Golgi apparatus and numerous mitochondria. In these cells the RER sometimes forms large intracisternal droplets (up to 7 m in diameter). Two of the animals exhibited a more uniform, high secretory activity. Large (about 2 m in diameter), pale vacuoles, probably of extracellular character, were found mostly in the vicinity of the perivascular septum. Their role in the release of MSH is discussed. The present data, which are discussed with reference to earlier findings (Forbes, 1972), form the morphological basis for an experimental study on regulation of MSH release (Larsson et al., 1979).Supported by grants from the Swedish Natural Science Research Council (to P. Meurling) and the Royal Physiographic Society of LundThe authors are indebted to Mrs. Ingrid Hallberg, Mrs. Kirsten Thörneby and Mrs. Lena Sandell for valuable technical assistance and to Miss Inger Norling for photographic aid  相似文献   

4.
Summary The anterior pituitaries from a series of female White-crowned Sparrows,Zonotrichia leucophrys pugetensis, in the periods of oviposition, incubation, and brooding under natural conditions, have been investigated by electron microscopy. The prolactin cells occur in cephalic lobe and are characterized by large (ca. 300–600 m), polymorphic electron-dense secretory granules and an extremely well developed, lamellated endoplasmic reticulum. During incubation and brooding it is only these prolactin cells that are in an activated secretory phase, as indicated by increase in number and size, extremely well developed endoplasmic reticulum, decrease in number of mature secretory granules, and by active formation of granules in the enlarged Golgi apparatus. In the late stages of brooding, and post-breeding, the prolactin cells regress with involution of the endoplasmic reticulum and Golgi apparatus, reaccumulation of granules, and the appearance of lysosomes.The gonadotropes of both the cephalic and caudal lobes undergo progressive morphologic changes through the course of the breeding period. They are numerous and active in the ovulating bird. They undergo gradual regression during the periods of incubation and brooding to become typical broody cells.This investigation was supported by Grant No. GF-33334, U.S.-Japan Cooperative Science Program of the National Science Foundation and by Grant No. GB-28080X, also from the National Science Foundation, to Professor Farner; and by Grant No. 5R040 Japan-U.S. Cooperative Science Program of Japan Association of Science Promotion, to Professor Mikami.  相似文献   

5.
Summary The ultrastructural immunocytochemical peroxidase-antiperoxidase technique used on serial sections reveals that certain corticotropic cells contain both ACTH and FSH, but not LH (ACTH/FSH cells). To determine the specificity of the anti-FSH staining in these cells, immunocytochemical absorption experiments were performed. The results indicate that (1) anti-FSH and -ACTH antisera are bound to different antigens in the corticotropic cell, and (2) anti-FSH staining is specific for a FSH-like antigen. In the ACTH/FSH cells both hormones are stored in the same secretory granules, distributed among other granules that contain only ACTH.  相似文献   

6.
The possible function of globules and irregular membrane-bound masses in the gonadotropin cells of the pituitary of Clarias gariepinus was studied. Strong secretory stimulation led to the disappearance of the secretory granules from gonadotropin cells but globules and irregular masses remained present. Acid phosphatase was detected enzyme-cytochemically in both globules and irregular masses. Radiolabelling with tritiated amino acids followed by autoradiography demonstrated that globules received radioactive material after secretory granules. The latter received radioactive material within 75 min of administration of radioactive amino acids but globules and irregular masses did not. Although some globules became radioactively labelled within 24 h of the administration of radioactive amino acids, irregular masses remained unlabelled during this period. Secretory granules reacted positively with antisera against and gonadotropin subunits, whereas globules and irregular masses only reacted with the antiserum against the subunit. A moderate anti-7B2 immunoreactivity was demonstrated in secretory granules and globules, whereas irregular masses labelled strongly. The combined cytological results indicate that globules and irregular masses are degradative, possibly crinophagic structures which develop by fusional events from secretory granules to globules and then to irregular masses.  相似文献   

7.
Summary The pancreatic islets of the guinea pig have been studied by light and electron microscopy. The B granules in glutaraldehyde-fixed tissue often are cup-shaped with an indentation visible on one side of the granule. Phosphotungstic acid hematoxylin (PTAH) positive cells have been characterized by electron microscopy as three subtypes based on the size of the secretory granules. Aa cells are the most common and have secretory granules around 200 m in diameter. Ab cells have large secretory granules around 300 m in diameter and are relatively infrequent. Ac cells are the least common and have small (160 m) granules. Characteristic D cells are identifiable by electron microscopy and, on the basis of the subsequent study (Munger, Caramia, and Lacy, 1965), are identified as the silver positive cells observed by light microscopy.This investigation was supported in part by United States Public Health Service research grants GM-10102 and GM-03784 from the Institute of General Medical Sciences, and AM-01226 from the Institute of Arthritis and Metabolic Diseases.-The authors wish to acknowledge the valuable technical assistance of Mrs. Aileen Sevier and Mrs. Lidia Donohue.  相似文献   

8.
Placing the clawed toad Xenopus laevis on a black background stimulates the melanotrope cells in the pars intermedia of the pituitary gland to release proopiomelanocortin (POMC)-derived peptides, including -MSH and N-acetyl-endorphin. In this study three types of secretory granules, electron-dense(130 nm Ø), moderately electron-dense ( 160 nm Ø) and electronlucent ( 180 nm Ø), have been identified in these cells. Apparently, only dark granules are formed by the Golgi apparatus and lucent granules release their contents via exocytosis. Immuno-electron microscopy (immunogold double labelling) of glutaraldehyde-fixed and freeze-substituted material shows that desacetyl--MSH and N-acetyl--endorphin coexist in all three granule types. Quantification of immunostaining revealed that immunoreactivities to these peptides are lowest in the dark granules and highest in the light ones. It is proposed that intragranular processing of POMC to immunoreactive desacetyl--MSH and N-acetyl--endorphin involves an increase in granule size and a decrease in granule electron density. Black background-induced activation of the melanotrope cell is reflected by an increase in immunoreactivity of the secretory granules to each of the antisera. This suggests that cell activation stimulates the formation of peptides by intragranular processing of POMC and/or of intermediate POMC-processing products. In addition, cell activation evoked an increase in the percentage of the granule population that reacts with anti-N-acetyl--endorphin, probably by stimulating intragranular acetylation of -endorphin. Apparently, this acetylation is a regulated event that occurs in the cytoplasm, independently from the acetylation of desacetyl--MSH which takes place near the plasmalemma at the time of granule exocytosis.  相似文献   

9.
Summary The fine structural characteristics of normal rat corticotrophs stained with anti-porcine ACTH1–39 serum were studied. At the ultrastructure level immunoreactive corticotrophs appear to comprise four distinct cell types: (1) large stellate cells (Siperstein cells) containing granules (170–250 nm in diameter) arranged in a peripheral row and usually embracing an acidophil; (2) elongate spindle-shaped cells (Moriarty cells) in which the secretory granules (170–250 nm in diameter) are distributed in a row or in small clusters in the peripheral cytoplasm; (3) oval or polygonal cells filled only with small secretory granules (130–170 nm in diameter), resembling the acidophil of small granules type (Yoshimura et al. 1974); and (4) polygonal or stellate cells filled with secretory granules of varying diameters (180–300 nm in diameter) and occasionally embracing an acidophil. The first type is the most common, but the others are infrequent. It is concluded that the criteria of Siperstein and Miller (1970) do not necessarily include all categories of rat corticotrophs.  相似文献   

10.
Summary The ultrastructure of the follicle gland and of some Gomori-positive neurosecretory cells in the lateral lobes of the cerebral ganglia of Lymnaea stagnalis is described.The follicle wall consists of epithelial cells containing secretion granules, and of processes of lateral lobe nerve cells bearing cilia, including the processes of bipolar neurosecretory B-cells. The ultrastructure of the follicle of Ancylus fluviatilis and Planorbarius corneus appeared to be very similar to that of L. stagnalis. From the ultrastructure of the follicle wall and lumen, no conclusions can be drawn as to its formerly proposed possible functions (sense organ, endocrine organ, neuroendocrine organ). Of three neurosecretory cell types (B-cells, canopy cells, droplet cells), the positivity to chrome-haematoxylin and paraldehyde-fuchsin appeared to be due to elementary granules of different size and appearance. This finding supports in a particular sense the view that results obtained with the neurosecretory stains, should be judged carefully. Also in ordinary neurons elementary granules can be found, at times in great numbers. Several types of elementary granules (at least 6) could be distinguished. Their character and function (neurohormones, neurotransmitters) are not clear.  相似文献   

11.
Summary Studies of possible neuroendocrine structures in the pulmonate gastropod Helisoma tenue show that cerebral fuchsinophilic neurons with electron-dense granules (mean diameter 1,500 Å) apparently release their secretory content in an intercerebral commissure neurohemal area near the mediodorsal bodies (MDB) or in the median labial nerve neurohemal area.The MDB cells have axon-like processes which branch and end at the cerebral surface, separated by a thin capsule from the cerebral fuchsinophilic cells. The presence of granules (mean diameter 800 Å) in the terminals of the MDB cells suggests cell body origin, distal transport and release of the granular secretory material. The secretory product may have an influence on cerebral fuchsinophilic neurons.Electron microscopy reveals the presence of granules of different sizes and densities in expanded neurites at the periphery of the intestinal nerve of the visceral ganglion which may indicate the presence of a neurohemal area. However, the granules in the intestinal nerve neurites and in the visceral ganglion fuchsinophilic cells are similar to granules found in the heart which also suggests that the granules may carry a neurotransmitter.This study was supported by NTH traineship USPHS T1 CA 5045 and NSF grant GB-6424X. The author would like to thank Professor Howard A. Bern for guidance and support during the course of this work, Mr. John Underhill and Mr. Albert Blaker for photographic assistance, and Mrs. Emily Reid for preparation of the diagram in Fig. 16.  相似文献   

12.
Summary Ultrastructural aspects of the secretory and the endocytotic pathways and the lysosomal system of corpus cardiacum glandular cells (CCG cells) of migratory locusts were studied using morphological, marker enzyme, immunocytochemical and tracer techniques. It is concluded that (1) the distribution of marker enzymes of trans Golgi cisternae and trans Golgi network (TGN) in locust CCG cells corresponds to that in most non-stimulated vertebrate secretory cell types; (2) the acid phosphatase-positive TGN in CCG cells is involved in sorting and packaging of secretory material and lysosomal enzymes; (3) these latter substances are produced continuously; (4) at the same time, superfluous secretory granules and other old cell organelles are degraded; (5) the remarkable endocytotic activity in the cell bodies and the minor endocytotic activity in cell processes are coupled mainly to constitutive uptake of nutritional and/or regulatory (macro)molecules, rather than to exocytosis; (6) plasma membrane recycling occurs mainly by direct fusion of tubular endosomal structures with the plasma membrane and little traffic passes the Golgi/TGN; and (7) so-called cytosomes arise mainly from autophagocytotic vacuoles and represent a special kind of complex secondary lysosomes involved in the final degradation of endogenous (cell organelles) and exogenous material.  相似文献   

13.
Summary After maternal fasting for 72 h the pancreatic cells of 18-day-old foetal rats show a conspicuous enrichment in secretory material, with an increase of pancreatic insulin concentration and a marked development of the rough endoplasmic reticulum and the Golgi apparatus.The morphometric analysis shows that the intracytoplasmic migration of the secretory granules is inhibited, principally inside the cell web. Consequently the number of secretory granules fused with plasma membrane decreases and this is associated with a decreased foetal plasma insulin.The difference in the ultrastructural aspect of the cells of foetuses from fasting mothers and of foetuses from fed mothers is less conspicuous at 19 days of gestation and progressively disappears at 20 and 21 days.The modifications in ultrastructural aspect and in functional state are discussed.  相似文献   

14.
Summary Twelve bovine adenohypophyses were prepared for light and electron microscopy of the cell types of pars distalis. Correlation between the light and electron microscopy was effected by use of alternate thin and thick sections. Cytological changes in the experimental animals were used as criteria for the identification of six different types of secretory cells.Two types of acidophils, alpha and epsilon cells, are recognized in peripheral area of the pars distalis by light and electron microscopy. The alpha cells contain orangeophilic secretory granules of a maximum diameter of 400–450 m and correspond to ordinary acidophils (STH cells). The second type, epsilon cells, contains larger, fuchsinophilic granules of 600 to 900 m in diameter, increase in number and granulation after pregnancy and thyroidectomy, and are thought to be prolactin cells (LTH cells).Two types of amphophils, zeta and delta 1 cells, were found in the central area of the pars distalis. The zeta cells contain smaller numbers of amphophilic, cored granules (200 m maximum diameter) and based on the comparison with literature on other species of animals, are designated as ACTH cells. The delta 1 cells are round or oval and contain very dense, spherical granules (250–300 m) which are stained red or reddish purple with PAS, aldehyde thionin and PAS-methyl blue methods. They show extreme enlargement and bizarre cytoplasmic appearance after castration and are designated tentatively as LH gonadotrophs or LH cells.Two types of basophils, beta and delta 2 cells, were also identified by correlative light and electron microscopy. The beta cells are polygonal in outline, distributed exclusively in the zona tuberalis and contain large, less dense secretory granules (300–400 m) which are stained selectively with Gomori's aldehyde fuchsin. After thyroidectomy, they lose their secretory granules and are transformed into large, vacuolated thyroidectomy cells. They are therefore, identified as thyrotrophs or TSH cells. The delta 2 cells are round, oval or polygonal in shape and contain basophilic granules ranging from 220 to 300 m in diameter. They show extreme enlargement and vacuolization due to the dilation of endoplasmic reticulum, after castration, and are designated tentatively as FSH gonadotrophs or FSH cells.The investigation reported herein was supported by a Scientific Research Grant (No. 291049) from the Ministry of Education of Japan.  相似文献   

15.
Summary An immunocytochemical technique using the peroxidase-antiperoxidase complex (PAP) was applied to identify and characterize the LH-secreting cells in the ovine pituitary at the ultrastructural level. These cells, round or oval in shape, possessing flattened cisternae of the rough endoplasmic reticulum, contain one class of secretory granules (mean diameter 250 nm) and large dense bodies (600 to 800 nm in diameter). LH molecules and the two subunits LH and LH were localized on the secretory granules and on the small granules near the Golgi complex. The large dense bodies, the cisternae of the endoplasmic reticulum and the saccules of the Golgi complex showed no reaction product.Abbreviations used in this Article O-LH ovine luteinizing hormone - b-LH bovine luteinizing hormone - p-LH porcine luteinizing hormone - p-LH porcine LH subunit - p-LH porcine LH subunit - O-FSH ovine follicle stimulating hormone - b-TSH bovine thyrotropic hormone - A-b LH antiserum to bovine LH - A-pLH antiserum to porcine LH subunit - A-pLH antiserum to porcine LH subunit  相似文献   

16.
Summary The dorsal tegument of the mature cercaria of Notocotylus attenuatus is a syncytial, cytoplasmic layer, containing two types of secretory granule which are identifiable ultrastructurally. The type 1 secretory bodies are electron lucid, whereas most type 2 granules have a banded appearance. The ventral tegument contains granules which are secreted from the type 3 cells; the type 3 granules are membrane bound, electron dense, and consist of both an amorphous and a finely striated zone. The type 4 cells mainly contain cigar-shaped granules consisting of an amorphous core surrounded by concentric striations. The granules exhibit structural variability in shape and content. The type 4 cells undergo a cellular migration to the tegument during encystment. The structure of the posterior-lateral glands and mode of secretion of the granules are described. Possible functions of microtubules are discussed for each cell type. Details of some secretory processes involved in the formation of the hemispherical cyst wall are described. The layers of the cyst wall may be related to the granular contents of the various parenchymal cells of the cercaria. The tegument of the metacercaria originates primarily from the cytoplasm of the type 1, type 2, type 3 and type 4 cells.  相似文献   

17.
Summary Although fluoride-labile protecting groups and linkers have been developed in solid-phase peptide synthesis to add an extra level of orthogonality, fluoride ions were found to convert -peptides to their corresponding - or -peptides in Glu(OtBu)- and Asp(OtBu)-containing peptidyl resins. Different peptide sequences and fluoride reagents were examined to determine the scope of this side reaction. CZE analysis was found to be a useful analytical technique to characterize peptides with respect to this phenomenon.  相似文献   

18.
Summary In order to revalue the effects of colchicine on incisor secretory ameloblasts, entire mice were perfused with Krebs solution supplemented with a buffer and amino acids, through the right common carotid artery. The normal ultrastructure of the cells was maintained for 2 h with the perfusate alone. When colchicine (0.3–3.0 g/ml) was added to the perfusate, it induced ultrastructural changes, such as the loss of cytoplasmic microtubules, the loss of secretory granules in Tomes' process, the abnormal accumulation and secretion of secretory granules, disarranged Golgi apparatus and the fragmentation of rough endoplasmic reticulum. Vesicles (150–400 nm in diameter) resembling immature secretory granules also accumulated, the degree of accumulation depending on the duration of colchicine treatment. The accumulation of secretory granules and these vesicles suggests that the intracellular transport system was affected by colchicine but that the production of secretory granules was continuous throughout the experimental period. The present perfusion system has enabled us to treat ameloblasts with an agent that is a useful experimental tool for elucidating cell functions, despite being lethal to animals in vivo.  相似文献   

19.
Summary The effects of synthetic thyrotropin-releasing hormone on pituitary prolactin and thyrotropic cells were investigated in adult male Rana perezi (formerly Rana ridibunda) frogs. Animals were given daily injections of synthetic thyrotropin-releasing hormone into the dorsal lymph sac. Prolactin and thyrotropic cells were identified by the colloidal-gold method, using anti-human prolactin and anti-human--thyrotropin hormone as primary antisera. The stereological parameters of the rough endoplasmic reticulum, Golgi complex, and secretory granules of prolactin and thyrotropic cells were evaluated by ultrastructural morphometry (point-counting method). Thyrotropin-releasing hormone caused cytological changes in both cell-types which were consistent with increased synthesis and release of both prolactin and thryrotropin. These changes were still significant after 48 h treatment in the case of thyrotropic cells, while in prolactin cells the thyrotropin-releasing hormone increased the number of secretory granules. After 6 days, the cells resembled essentially those used as controls. These results indicate that thyrotropin-releasing hormone stimulates the synthesis and release of prolactin and thyrotropin, and that the response of each cell type to this hypothalamic stimulus follows a different time-course.This work has been supported by grants no. 2184-83 and PB 86-0095 from the Comisión Interministerial para la Ciencia y Tecnología, Spain  相似文献   

20.
Zusammenfassung Die Mehrzahl der sezernierenden Zylinderzellen in den tubulösen Endstücken der Glandula bulbourethralis der Ziege sind Schleimzellen. Ihre großen, aufgehellten Sekretgranula, die fast den gesamten Zelleib ausfällen, liegen so dicht, daß sie sich gegenseitig abplatten. Einzelne von ihnen haben ihre Hüllmembran verloren und neigen zur Konfluenz. Alle Schleimkörnchen sind PAS-positiv, viele von ihnen zeigen eine neuraminidaselabile Alcianophilie bei pH 2,5. Neben den Schleimzellen findet man in der Glandula bulbourethralis der Ziege einen zweiten Zelltyp, der durch helle, blasenförmige Kerne, ein stark entfaltetes granuläres endoplasmatisches Retikulum mit dilatierten Zisternen sowie ein ausgedehntes supranukleares Golgi-Feld gekennzeichnet ist. Dieser zweite Zelltyp enthält sehr elektronendichte, isoliert liegende Sekretkugeln, welche lichtmikroskopisch eine Proteinreaktion geben. Zwischen beiden Zelltypen kommen morphologische Übergangsformen vor. Dies macht es wahrscheinlich, daß es sich bei den beiden Typen lediglich um Funktionsstadien einer einzigen Drüsenzelle handelt. Die sezernierende Oberfläche beider Zelltypen ist durch die Ausbildung interzellulärer Sekretkapillaren vergrößert. Diese sind bereits lichtmikroskopisch aufgrund ihrer kräftigen ATPase- und 5-Nucleotidaseaktivität zu identifizieren. Das gesamte Drüsenparenchym reagiert sehr stark auf unspezifische Esterase und deutlich positiv auf -D-Galactosidase, -D-Glucuronidase, Leucinaminopeptidase, Cytochromoxydase und Succinatdehydrogenase. Die letzten 5 Enzyme sind in den Schleimzellen in geringerer Konzentration festzustellen als in den Zellen mit Eiweißgranula.
Studies on the ultrastructure and histochemistry of the bulbourethral gland in the goat
Summary The majority of the secretory cells in the tubular endpieces of the caprine bulbourethral gland are mucous cells. Their closely packed, relatively large secretory granules exhibit a low electron density. Some granules have lost their limiting membrane, this results in the accumulation of irregularly outlined masses of secretory material. All mucous secretory granules are PAS-positive, many of them are characterized by an alcianophilia at pH 2.5 which is extinguished by pre-treatment with a neuraminidase solution. The second type of secretory bulbourethral cell exhibits light, spherical nuclei, a well developed rough endoplasmic reticulum with dilated cisterns and a large supranuclear Golgi-Complex. The cytoplasm contains smaller, highly electron dense granules which are — according to histochemical tests — of protein nature. The existence of transitional forms between both described cell types permits the conclusion that they must be regarded as functional stages of one common gland cell. The secretory surface of both cell types is increased by intercellular canaliculi which can be identified in the light microscope by their strong ATPase and 5-nucleotidase activities. The entire parenchyma of the gland is site of an exceptionally high esterase concentration. Furthermore, the gland cells contain considerable amounts of -D-galactosidase, -D-glucuronidase, leucine aminopeptidase, cytochrome oxydase and succinic dehydrogenase. These last five enzymes are histochemically more active in the protein secreting than in the mucus producing cell type.
Mit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.  相似文献   

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