MORPHOGENESIS OF CAPITATE GLANDULAR HAIRS OF CANNABIS SATIVA (CANNABACEAE)

The glandular secretory system in Cannabis sativa L. (marihuana) consists of three types of capitate glandular hairs (termed bulbous, capitate-sessile, and capitate-stalked) distinguishable by their morphology, development, and physiology. These gland types occur together in greatest abundance and developmental complexity on the abaxial surface of bracts which ensheath the developing ovary. Bulbous and capitate-sessile glands are initiated on very young bract primordia and attain maturity during early stages of bract growth. Capitate-stalked glands are initiated later in bract growth and undergo development and maturation on medium, to full sized bracts. Glands are epidermal in origin and derived, with one exception, from a single epidermal initial. The capitate-stalked gland is the exception and is of special interest because it possesses a multicellular stalk secondarily derived from surrounding epidermal and subepidermal cells. Glands differentiate early in development into an upper secretory portion and a subtending auxiliary portion. The secretory portion, depending on gland type, may range from a few cells to a large, flattened multicellular disc of secretory cells. The secretory portion produces a membrane-bound resinous product which caps the secretory cells. Capitate-stalked glands are considered to be of particular evolutionary significance because they may represent a gland type secondarily derived from existing capitate-sessile glands.     

TRICHOMES AND CANNABINOID CONTENT OF DEVELOPING LEAVES AND BRACTS OF CANNABIS SATIVA L. (CANNABACEAE)

Trichome density and type and cannabinoid content of leaves and bracts were quantitated during organ ontogeny for three clones of Cannabis sativa L. Trichome initiation and development were found to occur throughout leaf and bract ontogeny. On leaves, bulbous glands were more abundant than capitate-sessile glands for all clones, although differences in density for each gland type were evident between clones. On pistillate bracts, capitate-sessile glands were more abundant than the bulbous form on all clones, and both types decreased in relative density during bract ontogeny for each clone. The capitate-stalked gland, present on bracts but absent from vegetative leaves, increased in density during bract ontogeny. The capitate-stalked gland appeared to be initiated later than bulbous or capitate-sessile glands during bract development and on one clone it was first found midway in bract ontogeny. Nonglandular trichomes decreased in density during organ ontogeny, but the densities differed between leaves and bracts and also between clones. Specific regulatory mechanisms appear to exist to control the development of each trichome type independently. In addition, control of trichome density seems to be related to the plant organ and clone on which the gland type is located. Cannabinoid synthesis occurs throughout organ development and is selectively regulated in each organ. Typically, cannabinoid synthesis occurred at an increasing rate during bract development, whereas in developing leaves synthesis occurred at a decreasing rate. Cannabinoid content on a dry weight basis was generally greater for bracts than leaves. Analyses of leaves indicate that other tissues in addition to glands may contain cannabinoids, while for bracts the gland population can accommodate the cannabinoid content for this organ. The functional significance of trichomes and cannabinoids in relation to evolution is discussed.     

ULTRASTRUCTURAL DEVELOPMENT OF CAPITATE GLANDULAR HAIRS OF CANNABIS SATIVA L. (CANNABACEAE)

The capitate-sessile and capitate-stalked glands of the glandular secretory system in Cannabis, which are interpreted as lipophilic type glandular hairs, were studied from floral bracts of pistillate plants. These glands develop a flattened multicellular disc of secretory cells, which with the extruded secretory product forms the gland head and the auxiliary cells which support the gland head. The secretory product accumulates beneath a sheath derived from separation of the outer wall surface of the cellular disc. The ultrastructure of secretory cells in pre-secretory stages is characterized by a dense ground plasm, transitory lipid bodies and fibrillar material, and well developed endoplasmic reticulum. Dictyosomes and dictyosome-derived secretory vesicles are present, but never abundant. Secretory stages of gland development are characterized by abundant mitochondria and leucoplasts and by a large vacuolar system. Production of the secretory product is associated with plastids which increase in number and structural complexity. The plastids develop a paracrystalline body which nearly fills the mature plastid. Material interpreted as a secretion appears at the surface of plastids, migrates, and accumulates along the cell surface adjoining the secretory cavity. Extrusion of the material into the secretory cavity occurs directly through the plasma membrane-cell wall barrier.     

GLANDULAR STRUCTURES OF HOLOCARPHA AND THEIR ONTOGENY

Carlquist , Sherwin . (Rancho Santa Ana Botanic Garden, Claremont, California.) Glandular structures of Holocarpha and their ontogeny. Amer. Jour. Bot. 46(4): 300–308. Illus. 1959.—Two types of advanced glandular structures occur in the 4 species of the genus Holocarpha. Sessile disk-shaped glands occur at the tips of upper leaves and of involucral and receptacular bracts. Unlike all other glandular structures of Madinae which have been investigated, these originate from several protodermal initials rather than a single one. These glands, however, represent modifications of a glandular trichome. The other type of glandular structure, termed hollow-stalked trichome here, occurs on the outer surface of involucral and receptacular bracts. These trichomes originate from a single cell but differ from others in the formation of a hollow stalk, the wall of which is one cell in thickness. Mesophyll of the bract, often with an included vascular bundle, is present as an intrusion into the base of the hollow stalk. Corresponding to the advanced nature of the glandular structures, the leaves show specializations in the “inrolling” of margins. Upper leaves have a cylindrical organization of vascular tissue, whereas basal leaves are “normal” and leaves of the main stem are intermediate. The species of Holocarpha differ in certain details of leaf anatomy and structure of hollow-stalked trichomes. The systematic distribution of these is given. The essential unity of the various glandular structures of Madinae is discussed both in terms of mature structure and ontogeny, and the steps in the evolution of these are suggested.     

THE LEAF OF CALYCADENIA AND ITS GLANDULAR APPENDAGES

Carlquist , Sherwin . (Rancho Santa Ana Botanic Garden, Claremont, Calif.) The leaf of Calycadenia and its glandular appendages. Amer. Jour. Bot. 46(2) : 70-80. Illus. 1959.—Large tack-shaped glands are characteristic of the leaves of Calycadenia which are associated with the inflorescence. These glands may be divided into those which are terminal on leaves and those which occur laterally on the surface of the leaf. Lateral glands show stages early in their development which are identical with those of simpler trichomes of Madinae. Terminal glands, which possess more vascularization of the stalk, show a more modified form of development. Vascularization is not derived from protoderm, but from more deeply-seated cells. These cells are included in a zone of elongation which forms the stalk. Vascular bundles may extend to the base of glands which lack vascularization in their stalks. Tack-shaped glands are considered an advanced form of trichome in which internal tissues of the leaf are involved. Within the genus Calycadenia, ontogenetic and comparative studies suggest that the following characters are advanced: reduction to a single terminal gland, “inrolling” of margins to form a cylinder of bundles, concomitant with a central core of fibers or a pectic channel. Systematic distribution of gland occurrence and of types of foliar structure are given.     

Glandular trichomes of Humulus lupulus var. Brewer's Gold: Ontogeny and histochemical characterization of the secretion

Leaves of Humulus lupulus possess two types of glandular trichomes: - peltate (lupulin) and bulbous.
Peltate trichomes are formed from a protodermal cell by two anticlinal divisions in perpendicular planes, followed by two periclinal ones that give rise to the initials of the head cells, the basal and the stalk cells. Head cells divide successively in radial and irregular planes. Fully developed peltate trichomes are built of a glandular head consisting of 30 to 72 cells, four stalk cells and four basal cells.
Bulbous trichomes are also formed from a protodermal cell by an anticlinal division followed by two periclinal ones that produce the initials of the glandular head cells, and the basal and stalk cells. Fully developed bulbous trichomes consist of four (occasionally eight) head glandular cells, two stalk cells and two basal cells.
The density of peltate trichomes decreases with the expansion of the leaves.
Both peltate and bulbous trichomes secrete essential oils. Peltate trichomes are the preferential site for the synthesis of bitter resins. Tannic acids could not be detected histochemically either in peltate or in bulbous trichomes. Both types of trichomes produce secretion that accumulates in the subcuticular space, being released, in the case of bulbous trichomes, by rupture of the cuticle.     

The fine structure of epidermal glands of regenerating and mature globiferous pedicellariae of a sea urchin (Lytechinus pictus)

After a globiferous pedicellaria is lost from a sea urchin, a new appendage of the same kind is usually regenerated in the weeks that follow. During the latter part of regeneration, head glands and stalk glands, both of epidermal origin, develop from undifferentiated cells. Head gland cells begin morphological differentiation in the epidermis and then delaminate into the underlying dermis. In the formation of the stalk gland, by contrast, undifferentiated cells delaminate from the epidermis and then begin morphological differentiation in the dermis. During late regeneration, cells in the head and stalk glands are characterized by extensive rough endoplasmic reticulum distended with intracisternal material; moreover, the Golgi complex is closely associated with some of the large cytoplasmic vacuoles. The accumulating secretions of the two glands differ both in fine structure and in site of storage. Head gland secretions are stored intracellularly in the cytoplasmic vacuoles, while stalk gland secretions leave the gland cells in an apocrine fashion and are stored in an extracellular lumen. After regeneration, the mature cells of the head glands and stalk glands contain relatively little distended endoplasmic reticulum, although a Golgi complex is still present. Presumably, mature gland cells, in comparison to regenerating gland cells, produce relatively little secretion; instead, the glandular products elaborated during regeneration are probably stored in the mature glands with little augmentation or turnover.     

TRICHOMES OF CANNABIS SATIVA L. (CANNABACEAE)

The diversity of non-glandular and glandular hairs of Cannabis sativa L. (marihuana) are described by scanning electron microscopy. The non-glandular hairs are of two major types, as distinguished by size differences and locations, and all of them are highly silicified. The presence of silica as well as cystoliths of calcium carbonate help in the identification of marihuana even in its ash residues. X-ray microanalyses of Cannabis hairs are compared with those of Humulus lupulus and Lantana camera, whose hairs have been considered to resemble those of marihuana. Glandular hairs are found to be of two major categories. One group consists of glands whose heads are generally made up of eight cells and the other group whose heads are generally made up of two cells but never more than four cells. All glands of both categories are stalked. Some glands of the first category are massively stalked and these are restricted solely to anthers and bracts of staminate and pistillate plants. The massive stalk is considered to be made up of epidermal and hypodermal cells that have grown in response to some stimulation during anthesis. Fine details of the shoot system of Cannabis, such as cuticular ridges on epidermal cells, warty protuberances on non-glandular hairs, and surface views of glands in developing stages are also reported. Glandular hairs on the bracts of Humulus lupulus resemble those of Cannabis.     

Glandular Hair Formation in Origanum Species

Eleven Origanum species were investigated in this study. Inall these species, leaf glandular hairs have the same structureand they have been found to follow a common developmental pattern.They originate from a single protodermal cell which dividessuccessively parallel to leaf surface to give rise to the footcell, the stalk cell and the mother cell of the head. The latterundergoes a series of symmetric and asymmetric anticlinal divisionsresulting in a 12-celled head (four small cells in the centreand eight large cells peripherally arranged). Determinationof the glandular scale morphology within the Lamiaceae generaand further comparison between the various gland types wouldbe helpful to systematics. Origanum species, Lamiaceae, glandular hair, development     

ULTRASTRUCTURE OF GLANDULAR TRICHOMES OF LEAVES OF NICOTIANA TABACUM L., cv XANTHI

Electron microscopy confirms previous light microscope observations that tobacco leaf trichomes are glandular and that there are two different types. Both the tall trichome (multicellular stalk, unicellular or multicellular head) and the short trichome (unicellular stalk; multicellular head) exhibit characteristics common to gland cells—a dense cytoplasm, numerous mitochondria, and little vacuolation. The tall trichome contains structurally well developed chloroplasts and an elaborate network of endoplasmic reticulum. The short trichome contains undifferentiated plastids and endoplasmic reticulum which parallels the nucleus and plasmalemma. Few dictyosomes are seen either in the short trichome or in the tall trichome. The short trichome appears to undergo structural changes concurrently with the appearance of secretory product within the cells. The most noticeable change is the formation of the extraplasmic space between the cell wall and the plasmalemma. Electron dense secretory product is observed between the plasmalemma and the cell wall and within the intercellular spaces.     

野薄荷叶腺毛的发育形态学研究

研究了分布于我国东北部的野薄荷叶表面腺毛的结构和发育。主要腺毛有两种：多细胞盾状腺毛和弯曲单细胞头状腺毛。多细胞盾状腺毛由１个基细胞,１个柄细胞和一个由８－１２个分泌细胞构成的头部组成。单细胞头状腺毛由１个基细胞,１个柄细胞和一个分泌细胞头部构成。     

日本血吸虫尾蚴的组织化学及扫描电镜观察

日本血吸虫尾蚴具有一个头腺、2对前钻腺及3对后钻腺,其解剖学位置、构造、化学成分及功能均有区别。钻腺分泌物含有多糖酶及蛋白酶。尾蚴的前端系特化的头器结构,它具有半月形嵴、钻腺开口及乳突。     

GLAND DISTRIBUTION AND CANNABINOID CONTENT IN CLONES OF CANNABIS SATIVA L.

The relationship between glandular trichomes and cannabinoid content in Cannabis sativa L. was investigated. Three strains of Cannabis, which are annuals, were selected for either a drug, a non-drug, or a fiber trait and then cloned to provide genetically uniform material for analyses over several years. The distribution of the number and type of glands was determined for several organs of different ages including the bract and its subtending monoleaflet leaf and the compound leaf on pistillate plants. Quantitation of glands on these structures was integrated with gas chromatographic analyses of organ cannabinoid profiles. A negative correlation was found between cannabinoid content and gland number for each of the three clones. Isolated heads of the capitate-stalked glands also were analyzed for cannabinoid content and found to vary in relation to clone and gland age. These studies indicate that cannabinoids may occur in plant cells other than glandular trichomes. The results of these studies emphasize the need for stringent sampling procedures in micromorphological studies on trichome distribution and analytical determinations of cannabinoid content in Cannabis.     

The architecture of the accessory reproductive glands of the male desert locust: 1: types of glands and their secretions

The accessory reproductive glands of the male desert locust were studied by histological, histochemical, and phase-contrast techniques. It was found that the characteristics of the glandular epithelium and their corresponding secretions permit the division of the accessory glands into nine distinct types. Three types produce coarsely granular mucopolysaccharide secretions (glands 1, 11, and 12); three types produce finely fibrous mucopolysaccharide secretions (glands 2, 4, and 7-10, 13-15); one type produces a globular mucopolysaccharide or mucoproteinaceous secretion (gland 6); one type produces an acidic lipoprotein complex (glands 3 and 5); and one is the functional seminal vesicle (gland 16). Consequently, the various secretions are separated as a result of a vertical segregatign of the various cell types that are responsible for glandular activity.     

甘草腺毛的形态发生和组织化学研究

利用扫描电镜及薄切片技术对甘草的腺毛形态发生和发育过程进行了观察,并对腺毛发育过程中黄酮类成分积累进行了组织化学定位研究。结果表明：甘草腺毛为多细胞构成的盾状腺毛,有长柄和短柄2种类型;前者主要分布在花萼片上,而后者主要分布于叶片上。组化鉴定结果显示：腺毛中存在着黄酮类成分、其他亲脂类和非纤维素多糖类成分;在腺毛的发育过程中,黄酮类物质是随腺毛的发育成熟,在头部盘状结构的分泌细胞及角质层下腔中积累。研究结果对进一步探讨甘草叶中黄酮类成分的合成及其作用提供科学依据。     

Development of peltate glandular trichomes of peppermint

Cryofixation and conventional chemical fixation methods were employed to examine the ultrastructure of developing peltate glandular trichomes of peppermint (Mentha x piperita). Our results are discussed in relation to monoterpene production and the mechanism of essential oil secretion. Peltate glands arise as epidermal protuberances (initials) that divide asymmetrically to produce a vacuolate basal cell, a stalk cell, and a cytoplasmically dense apical cell. Further divisions of the apical cell produce a peltate trichome with one basal cell, one stalk cell, and eight glandular (secretory) disc cells. Presecretory gland cells resemble meristematic cells because they contain proplastids, small vacuoles, and large nuclei. The secretory phase coincides with the separation and filling of the sub-cuticular oil storage space, the maturation of glandular disc cell leucoplasts in which monoterpene biosynthesis is known to be initiated, and the formation of extensive smooth endoplasmic reticulum at which hydroxylation steps of the monoterpene biosynthetic pathway occur. The smooth endoplasmic reticulum of the secretory cells appears to form associations with both the leucoplasts and the plasma membrane bordering the sub-cuticular oil storage cavity, often contains densely staining material, and may be involved with the transport of the monoterpene-rich secretion product. Associated changes in the ultrastructure of the secretory stage stalk cell are also described, as is the ultrastructure of the fragile post-secretory gland for which cryofixation methods are particularly well suited for the preservation of organizational integrity.     

Development and ultrastructure of glandular trichomes in<Emphasis Type="Italic">pelargonium x fragrans</Emphasis> ‘mabel grey’ (geraniaceae)

The glandular trichomes of leaves fromPelargonium xfragrans ‘Mabel Grey’ (Geraniaceae) were examined by light, scanning, and transmission electron microscopy. These trichomes had unicellular globular heads and stalks of different lengths and features. Two types were classified: Type I, with an elongated, large head and a short (100 μm), cylindrical stalk that was more apparent on the adaxial surface; and Type II, with a spherical, small head and a long (300μm), conical stalk that was more pronounced on the abaxial surface. The ultrastructure of secretory cells from both types was distinguished by a well-developed endoplasmic reticulum, mitochondria, plastids, dictyosomes, and numerous vacuoles that likely were involved in the storage and transport of lipophilic substances. Plasmodesmata were frequent on the walls of the secretory and stalked cells. Here, we discuss the implication of structural differentiation in these trichomes.     

紫苏腺毛的形态发生研究

紫苏叶上有两种腺毛：质状腺毛和头状腺毛。两者都具１个基细胞、１个柄细胞和头部。前者的头部可由１、２、４或８个分泌细胞组成,扩展成质状;后者的头部由１、２或４个分泌细胞组成,聚成圆球状。两种腺毛的原始细胞都来源于原表皮细胞,经两次平周分裂产生基细胞、柄细胞和顶细胞。在腺毛后期的形态发生中,柄细胞的分化状态决定腺毛的类型。若柄细胞保持扁平关且处于分生状态时,其顶细胞将发育成质状腺毛的头部;若柄细胞纵向     

紫苏腺毛的形态发生研究

紫苏叶上有两种腺毛:盾状腺毛和头状腺毛。两者都具1个基细胞、1个柄细胞和头部。前者的头部可由1、2、4或8个分泌细胞组成,扩展成盾状;后者的头部由1、2或4个分泌细胞组成,聚成圆球状。两种腺毛的原始细胞都来源于原表皮细胞,经两次平周分裂产生基细胞、柄细胞和顶细胞。在腺毛后期的形态发生中,柄细胞的分化状态决定腺毛的类型。若柄细胞保持扁平状且处于分生状态时,其顶细胞将发育成盾状腺毛的头部;若柄细胞纵向引长并迅速液泡化时,其顶细胞将发育成头状腺毛的头部。     

Salivary glands and salivary pumps in adult Nymphalidae (Lepidoptera)

The salivary glands and salivary pumps were investigated by means of dissection and serial semithin sections in order to expose the anatomy and histology of Nymphalidae in relation to feeding ecology. The paired salivary glands are tubular, they begin in the head, and extend through the thorax into the abdomen. The epithelium is a unicellular layer consisting of a single cell type. Despite the uniform composition, each salivary gland can be divided into five anatomically and histologically distinct regions. The bulbous end region of the gland lies within the abdomen and is composed of highly prismatic glandular cells with large vacuoles in their cell bodies. The tubular secretion region extends into the thorax where it forms large loops running backward and forward. It is composed of glandular cells that lack large vacuoles. The salivary duct lies in the thorax and also shows a looped formation but is composed of flat epithelial cells. The salivary reservoir begins in the prothorax and reaches the head. Its cells are hemispherical and bulge out into the large lumen of the tube. In the head the outlet tube connects the left and right halves of the salivary gland, and its epithelial cells are flat. The salivary pump lies in the head ventral to the sucking pump and leads directly into the food canal of the proboscis. It is not part of the salivary gland but is derived from the salivarium. Both the thin cuticle of the roof of the salivary pump and the thick bottom are ventrally arched. Paired muscles extend from the hypopharyngeal ridges and obviously serve as dilators for the pump. A functional interpretation of the salivary pump suggests that when not in use, the dilators are not contracted and the pump is tightly closed due to its own elasticity. When the dilator muscles repeatedly contract, the saliva is forced forward into the food canal of the proboscis. The salivary gland anatomy was found to be similar to other Lepidoptera. Furthermore, the histology of the salivary glands is identical in all examined butterflies, even in the species which exhibit specialized pollen-feeding behavior.