芸香科植物茎分泌囊的研究

利用石蜡切片和薄切片方法对芸香料１４属２３种和１变种植物幼茎中分泌囊的分布和结构进行了比较研究。在芸香科植物茎中,靠近表皮的皮层中分布一轮分泌囊,分泌均由鞘细胞和一层上皮细胞围绕圆形腔隙构成,上皮细胞扁平,细胞壁薄,完整。鞘细胞１－５层。在不同亚科、不同属之间,分泌囊的差异仅仅表现在分泌腔的大小和鞘细胞的层数方面。一般草本类型属植物     

中国芸香科植物叶分泌囊比较解剖学研究

利用整体透明、石腊和薄切片方法对芸香科22属,40种和2变种植物叶分泌囊的形态结构和分 布进行了比较研究。成熟分泌囊都由鞘细胞和一层上皮细胞围绕圆形腔隙构成,上皮细胞扁平,细胞壁 薄、完整,故分泌囊属裂生方式发生。鞘细胞1～5层,不同种类的层数有变化,个别种缺乏。内层鞘细 胞为扁平的薄壁细胞,外层的细胞壁较厚。分泌囊的形态结构、着生位置和分布密度等在不同属或不同 种间存在一定差异。根据分泌囊在叶中的分布位置和形态结构特点,可将其划分为:叶缘齿缝分泌囊, 叶肉分泌囊和两者混合型。叶肉分泌囊又可分海绵组织分泌囊和栅栏组织分泌囊。在此基础上对该科各类型分泌囊的形态演化关系以及各亚科或各属间的亲缘关系进行了探讨。     

Secretory Cavity Development and Its Relationship with the Accumulation of Essential Oil in Fruits of Citrus medica L. var. sarcodactylis （Noot.） Swingle

The developmental types of secretory cavities in Citrus remain controversial. The relationship between secretory cavity development and the accumulation of essential oil in fruits of Citrus species is also unknown. In order to develop better insights into these problems, histological, histochemical, and cytochemical methods were used to investigate secretory cavity development and the accumulation of essential oll at different developmental stages of fruits of Citrus medica L. var. sarcodactylis （Noot.） Swingle. The results indicate that the secretory cavity of the variety seemed to originate from an epidermal cell and a subepldermal cell. These two cells underwent successive divisions, resulting In the formation of two parts： （Ⅰ） a conical cap; and （Ⅱ） a globular gland. The formation of the lumen was schlzolysigenous. Regular changes in the size of vacuoles and the accumulation of essential oil were revealed during the process of secretory cavity development. In addition, when fruits were a light yellow or golden color, the structure of secretory cavities was well developed and the content of essential oil in a single fruit reached a maximum. It would be most appropriate to collect the fruit as a medicinal material at this time.     

COMPARATIVE DEVELOPMENT OF SECRETORY CAVITIES IN THE TRIBES AMORPHEAE AND PSORALEEAE (LEGUMINOSAE: PAPILIONOIDEAE)

The tribes Amorpheae and Psoraleeae of the Leguminosae (Papilionoidae) share the characteristics of one-seeded fruits and gland-dotted foliage. Because of this, they traditionally have been considered closely related (either a single tribe or two closely related tribes). However, Barneby (1977) has suggested that the Amorpheae and Psoraleeae are not close but previously had been combined on the basis of a superficial resemblance. This paper describes the structure of the secretory cavities responsible for the gland dots. Approximately 50% of the species of each tribe were surveyed for cavity structure with leaflet clearings. Eight species were then chosen for developmental studies of their glands. Several distinct kinds of secretory cavities are present in these plants. Trabeculate cavities (found only in the Psoraleeae) are traversed by many elongated cells. This type of cavity and nontrabeculate cavities of the Psoraleeae initiate with localized dorsiventral elongation of protodermal cells to form a hemispherical protuberance on the leaf primordium surface. Development proceeds with separation of the cells of a protuberance along their lateral walls facing the protuberance center. As the leaf expands, the protuberance sinks until its apex is flush with the leaf surface. The result is a cavity lined by an epithelium of modified epidermal cells. Trabeculate cavities have more cells in the initial protodermal bump than nontrabeculate “epidermal” cavities, and the central cells of the protuberance are not involved in epithelium formation, but become separated from other cells on all lateral sides, transversing the cavity as trabeculae. Cavities of the Amorpheae are all nontrabeculate and subepidermal. They initiate with periclinal divisions of protodermal cells that result in two cell layers. The exterior layer differentiates into epidermis, while the interior layer divides to produce a small spherical group of cells (“epithelial initials”). Schizogeny occurs in the center of these cells to produce an epithelium-lined cavity. Previous studies of cavity development in the Amorpheae described lysigenous and schizo-lysigenous cavities for most species. These early reports are reviewed, and the possible role of preparation artifacts in producing images of lysigenous development in general is discussed.     

Studles on the Initiation and Development of Gutta-containing Cells of Eucommia ulmoides Oliv.

The gutta-containing structure in the cortex of a stem of Eucommia ulmoides is a filamentous, secretory cell. Observation of thin sections revealed that when the procambium of a stem differentiated into the earliest sieve elenents of protophloem, the initials cells appeared as daughter cells originated from a longitudinally equational or unequational divisions, or from the terminal cell formed by several transverse divisions of some cortical ground meristematic cells. Before the ceils of the cortical ground'meristem ceased to divide, the initial cells, in an arbitrary position of origin, develops continuously. These initial cells were able to be distinguished from the surrounding cells by their large length/width ratio, the presence of elliptical nuclei and dense cytoplasm, etc. Later, both ends of the initial cells extended rapidly through intrusive growth, forming the very long, thin and filamentous unicell with two dilated terminations. During development, the gutta particles were gradually synthesized and accumulated in the cytoplasm, where as the organelles degenerated progressively. In the muture gutta-containing cell, the cell cavity was filled with gutta particles, the nucleus as well as other organelles was disintegrated leaving an intact fibrous cell wall.     

Programmed cell death of secretory cavity cells in fruits of Citrus grandis cv. Tomentosa is associated with activation of caspase 3-like protease

Previously, we found that secretory cell degradation typically occurred through programmed cell death during secretory cavity development in Citrus sinensis L. (Osbeck). This finding indicated that secretory cavities could be utilized as a new cell biology model for investigating the regulatory mechanisms of plant programmed cell death. To study further the programmed cell death during secretory cavity development in Citrus fruit, we studied the morphogenetic characteristics of secretory cavities during their development in Citrus grandis cv. Tomentosa. Using light microscope- and electron microscope-TUNEL assays, immunohistochemistry and immunocytochemistry, we described the precise spatial and temporal alterations in caspase 3-like distribution, chromatin condensation and DNA fragmentation during the programmed cell death of secretory cavity cells. Caspase 3-like was found to be significantly located in both the cytoplasm and the nucleus of secretory cavity cells undergoing programmed cell death, and caspase 3-like is closely associated with chromatin condensation and DNA fragmentation. Interestingly, both caspase 3-like and DNA fragmentation were detected in the nucleoli. Our findings suggest that caspase 3-like may be involved in the programmed cell death of secretory cavity cells, especially in chromatin condensation, DNA fragmentation, nuclear degradation and the degradation of certain organelles.     

贯叶连翘分泌结构的发育及其内含物积累的研究

应用半薄切片和组织化学观察结果表明,贯叶连翘植物内具有分泌囊和分泌细胞团,叶片,萼片,花瓣中都存在2种分泌结构,分泌囊尚存在于果壁内,而分泌细胞团还在花药和茎棱中 ,2种分泌结构均起源于幼叶的基本分生组织内,其原始细胞形态相似,都垂周分裂为2子细胞,接着分别以不同分裂方式形成原始细胞群,并分化出鞘细胞,以后分 囊原始细胞群以裂生方式发育成由1层鞘细胞,1层上皮细胞包围着分泌腔组成的分泌囊,而分泌细胞团原始细胞群则继续增加细胞数目和体积发育成2－4层鞘细胞包围紧密排列的分泌细胞组成的分泌细胞团,其中给终未出现分泌腔,组织[化学试验表明,前者产生和贮存油类物质,而后者产生和贮存金丝桃素类物质。     

Structure and development of the secretory cavities of Myrtus communis leaves

The structure and development of Myrtus communis L. secretory cavities has been studied in young and expanded leaves, using light and scanning electron microscope. Secretory cavities are continuously formed during leaf development, but in mature leaves the rhythm of their appearance shows steep decrease. Each secretory cavity is developed from a single epidermal cell, which undergoes a periclinal division followed by anticlinal and several oblique cell divisions. The lumen of the secretory cavity is initiated by cell wall separation, i.e., schizogenously. The secretory cells line the cavity, where the secreted material is collected. Secretory cavities are covered by modified epidermal cells, which do not seem to form any special aperture. Essential oils seem to be discharged after mechanical treatment of the leaf.     

Observations on the development of the foliar secretory cavities of Porophyllum lanceolatum (Asteraceae)

The lipophilic secretory cavities observed in the leaf of Porophyllum lanceolatum (Asteraceae) are scattered throughout the lamina and around its crenate margins. In the young leaf the cavities are initiated, and their development completed, while the surrounding tissues are still at early stages of differentiation. The cavity lumen has a lysigenous origin. Cell lysis, expansion of the developing leaf and, probably, the pressure exerted by the accumulation of secretory products, are believed to account for the gradual enlargement of the lumen. Concomitantly with ctll disintegration, which occurs throughout development, divisions take place in all cells of the gland. A mature cavity has a multilayered epithelium. Histochemical tests for RNA, proteins, phenolics and pectic polysaccharides revealed intense staining of the content of the epithelial cells in the early stages of cavity development, and a decrease in staining towards its maturity. Staining for lipids is intense in all developmental stages. Tests on the material observed in the lumen of mature cavities, show positive results for lipids, pectic polysaccharides and phenolics.     

The Relationship Between Oil Gland and Fruit Development in Washington Navel Orange (Citrus sinensis L. Osbeck)

Changes in structure, size and number of oil glands locatedin the fruit rind were assessed in developing fruit of the WashingtonNavel orange (Citrus sinensis L. Osbeck) from pre-anthesis tofruit maturity. Initiation of oil glands was found to be restrictedto early fruit development. Glands continued to develop throughoutfruit growth, until all reached maturity by a fruit size of30 to 50 mm diameter. Mature glands continued to enlarge withfruit growth. Mature fruit had between 8 000 and 12 000 oilglands. Anatomical studies of the fruit rind were carried outusing light microscopy on samples prepared by different tissueprocessing methods. Glands were found to develop from a clusterof cells adjacent to the fruit epidermis, into a structure consistingof a central cavity surrounded by several layers of epithelialcells. All glands were joined to the fruit epidermis, irrespectiveof their stage of development. Neither lignin nor suberin waspresent in the gland. Gland cavity formation appeared to involveschizogeny. Copyright 2001 Annals of Botany Company Washington Navel orange, Citrus sinensis L. Osbeck, fruit development, secretory cavity, oil gland, image analysis, light microscopy     

Structural and ultrastructural aspects of ontogenesis and differentiation of resin secretory cavities in Hymenaea stigonocarpa (Fabaceae‐Caesalpinioideae) leaves

Cell lysis in the formation of secretory cavities in plants has been questioned by some authors and considered as result of technical artifacts. To describe the formation of secretory resin cavities in Hymenaea stigonocarpa leaves, leaflet samples at different stages of differentiation were collected, fixed, and processed for light and electron microscopy as per usual methods. The initial cells of secretory resin cavities are protodermal and grow towards the mesophyll ground meristem; these cells then divide producing cell groups that are distinguished by the shape and arrangement of cytoplasm, and density. At the initial stages of differentiation of the secretory cavities, some central cells in these groups show dark cytoplasm and condensed nuclear chromatin. Later, there is cell wall loosening, tonoplast and plasmalemma rupture resulting in cell death. These cells, however, maintain organelle integrity until lysis, when the cell wall degrades and the plasmalemma ruptures, releasing protoplast residues, marked characteristics of programmed cell death. The secretory epithelium remains active until complete leaf expansion when the cavity is filled with resin and the secretory activity ceases. There are no wall residues between central cells in adult cavities. Our results demonstrate lysigeny and the importance of ontogenetic studies in determining the origin of secretory cavities.     

Cell Degeneration and Cavity Formation in the Endocrine Pancreas of the Hagfish Myxine glutinosa

Abstract Cell degeneration within the islet lobules of the pancreas of Myxine glutinosa appears to be a natural feature of the gland in this species. Degenerating cells bordering a lobule cavity discharge debris directly into the cavity, while cells undergoing degeneration at sites distant from such cavities discharge debris into the surrounding intercellular space. The latter process may well act as a locus for local degeneration within the lobule and the formation of a lobule cavity.     

FINE STRUCTURE OF EPITHELIAL CANAL CELLS IN PETIOLES OF XANTHIUM PENSYLVANICUM

Secretory canals, lined with an epithelium, occur in many families, e.g., Umbelliferae, Compositae. These canals are said to extend continuously through the root and shoot systems and are known, in some cases, to secrete resins, essential oils, etc. In Xanthium the canals arise schizogenously from cells derived from canal initials. Subsequent divisions lead to a ring of 7–12 epithelial cells surrounding a central cavity. During maturation the epithelium becomes crushed and obliterated. Canals were examined in petioles of Xanthium pensyhanicum (Cocklebur) grown under long day illumination to maintain vegetative growth. The fine structure of the canal and its epithelium was studied by electron microscopy of thin sections cut transverse to the principal axis of petioles from leaves in an early stage of development. The canal proper is delimited by walls of epithelial cells which protrude into a scallop shaped cavity. In comparison to the surrounding parenchyma, the epithelial cells are smaller, cytoplasmically more dense, and less vacuolate. The epithelium contains pleomorphic starch-free plastids with planar thylakoids frequently stacked into grana; thus, the plastids are presumed photosynthetically active. Mitochondria are abundant and often dense. The cytoplasm is rich in free polysomes, and smooth endoplasmic reticulum predominates over the rough form. Spheroidal granules averaging about 530 nm in diameter are numerous in the epithelium and appear at lower concentration in neighboring cells. Many features of fine structure of the epithelial cells suggest that a high metabolic activity is present in this tissue during this early stage of development. A possible function of the canals is defense against insect predation and animal grazing.     

Morphological characteristics of sclerotia formed from hyphae of <Emphasis Type="Italic">Grifola umbellata</Emphasis> under artificial conditions

The morphological characteristics of sclerotia were induced in cultures of the fungus Grifola umbellata by introducing an unidentified companion fungus were studied by light microscope, scanning and transmission electron microscope (SEM and TEM). Light microscope and SEM investigations of developing sclerotia revealed that aerial mycelial hyphae diminished with age, and mature sclerotia had two tissue layers, the rind and medulla. The medulla was comprised of thin and thick-walled hyphae of varying diameter. The thick-walled cells always formed below the hyphal tips. Retraction of the cytoplasm was accompanied by the thickening of cell wall. There were crystalline initials in the newly formed sclerotium. Crystalline initials were always formed in the tip of medullary hyphae and were not of regular shape. A series of changes occurred in the cells in which the crystalline initials would be formed, such as enlargement of size, formation of one or several large vacuoles. Crystalline initials developed via amorphous materials in the cytoplasm deposited in the vacuoles. At last crystalline initials was released by degradation of the cell wall.     

SECRETORY VESICLE FORMATION IN GLANDULAR TRICHOMES OF CANNABIS SATIVA (CANNABACEAE)

Formation of secretory vesicles in the noncellular secretory cavity of glandular trichomes of Cannabis saliva L. was examined by transmission electron microscopy. Two patterns of vesicle formation occurred during gland morphogenesis. 1) During initial phases of cavity formation small hyaline areas arose in the wall near the plasma membrane of the disc cell. Hyaline areas of elongated shape and different sizes were distributed throughout the wall and adjacent to the secretory cavity. Hyaline areas increased in size, some possibly fusing with others. These hyaline areas, possessing a membrane, moved into the cavity where they formed vesicles. As membraned vesicles they developed a more or less round shape and their contents became electron-dense. 2) During development of the secretory cavity and when abundant secretions were present in the disc cells, these secretions passed through the wall to accumulate as membraned vesicles of different sizes in the cavity. As secretions emerged from the wall, a membrane of wall origin delimited the secretory material from cavity contents. Vesicles released from the wall migrated in the secretory cavity and contacted the sheath where their contents permeated into the subcuticular wall as large or diffused quantities of secretions. In the subcuticular wall these secretions migrated to the wall–cuticle interface where they contributed to structural thickening of the cuticle. This study demonstrates that the secretory process in glands of Cannabis involves not only secretion of materials from the disc cell, but that the disc cell somehow packages these secretions into membraned vesicles outside the cell wall prior to deposition into the secretory cavity for subsequent structural development of the sheath.     

吴茱萸果实中分泌囊的发生和发育研究

吴茱萸果皮内分布有许多分泌囊。我们作了发育解剖学方面的研究。在花蕾期,雌蕊的子房中分泌囊原始细胞即开始发生,它起源于单个表皮细胞和其内的1—4层薄壁细胞。分泌囊最初为裂生,后期由于上皮细胞的破毁,其腔隙逐渐扩大,因此,腔隙发生方式应属裂溶生型。成熟分泌囊是由多层鞘细胞和上皮细胞包围圆形腔隙构成。     

Developmental Ultrastructure of Glandular Trichomes of <Emphasis Type="Italic">Rosmarinus officinalis</Emphasis>: Secretory Cavity and Secretory Vesicle Formation

Glandular trichomes in the leaf lamina of Rosmarinus officinalis L. were examined by scanning and transmission electron microscopy. The leaves were characterized by an abundance of two types of glandular trichomes—small capitate and large peltate glandular trichomes. In addition to the glandular trichomes, numerous non-glandular trichomes were present on the abaxial surface of the leaf. These trichomes mainly predominated on the midrib, whereas glandular trichomes occurred on non-vein areas. At the initial phase of secretory cavity formation, hyaline areas were abundant in periclinal walls of head cells, while they were not observed in the anticlinal walls. The hyaline areas gradually increased in size, fusing with other areas throughout the wall. Loose wall material adjacent to hyaline areas was released from the head cell walls and migrated into the secretory cavities. As the secretory cavities continued to enlarge, the new vesicles emerging into the secretory cavities from the walls of head cells became surrounded with the surface of a typical membrane. They developed a round shape, but the contours of the vesicle surfaces appeared polygonal when tightly packed inside a cavity. These vesicles varied in size; small vesicles often possessed electron-dense contents, while large vesicles contained electron-light contents.     

DEVELOPMENTAL ANATOMY OF SECRETORY CAVITIES IN THE MICROSPOROPHYLLS OF GINKGO BILOBA L.

Two to five secretory cavities develop in the hump region of the microsporophylls of Ginkgo biloba. A developing cavity is first recognized as a spherical pocket of large, densely cytoplasmic cells (central secretory cells) in the median portion of a microsporophyll primordium. These cells degenerate and a small cavity is formed which is filled with the contents of the degenerating cells. Flattened incurved cells (parietal secretory cells) develop around the disintegrating central secretory cells and slough off into the enlarging cavity. Thus, the cavities develop by lysigeny. A mature cavity is surrounded by senescent parietal secretory cells, scanty parenchyma, and a loosely fitting epidermis. Histochemical tests indicate the presence of lipid and pectic substances in the cavities. Previous reports on the morphological interpretation and possible function role of the cavities are discussed in the light of the present investigation.     

Ultrastructural characteristics of the salivary glands of the taiga tick, Ixodes persulcatus (Ixodidae). I. The granulosecreting alveoles of the fasting female

Two types of granulosecreting alveoles were found in salivary glands of hungry females by means of electron microscopy of ultrafine sections. Alveoles of the IInd type occur in the anterior helf of the gland. They are not numerous and consist of three types of secretory cells (A, B, C) surrounding the inneralveolar cavity. The secretory cells are separated from each other and from the basal membrane by the strands of the epithelial cells P. Three types of spherical inclusions were found in the secretory cells. They differ in size, electron density and intensity of staining of half-fine sections with toluidin blue. The apical cytoplasmatic membrane of secretory cells bears numerous microvilli. Alveoles of the IIIrd type, which constitute the main mass of the gland tissue, have a narrow slit-like inneralveolar cavity. The basal part of the alveole is formed by 3--4 large cells filled with large spherical electron-transparent vacuoles of the secretion. The apical part of the alveole is occupied by 9 to 11 cells E, whose cytoplasm is filled with numerous flat cisternae of granular endoplasmatic reticulum and small and medium secretory vacuoles of different electron density. Alveoles of the IInd and IIIrd type of I. persulcatus are not identical with those of Hyalomma asiaticum, Boophilus microplus and other members of the subfamily Amblyomminae.