STUDIES ON THE ONTOGENY OF THE DWARF MISTLETOES,ARCEUTHOBIUM. II. HOMOLOGY OF THE ENDOPHYTIC SYSTEM

Cohen , L. I. (Washington State U., Pullman.) Studies on the ontogeny of the dwarf mistletoes, Arceuthobium. H. Homology of the endophytic system. Amor. Jour. Bot. 50(5): 409–417. Illus. 1963.—Development of the seedling in Arceuthobium, as well as the mode of penetration and infection, is described. The promeristem of the root apex of the seedling, as in the embryo, is composed of 4 zones: (1) a uniseriate layer of apical surface initials; (2) a subapical zone of central initials; (3) a peripheral zone; and (4) a zone of procambial initials. After germination, the seedling grows on the host surface until the root apex is confronted by a spur shoot or by a fragment of raised bark. The root apex becomes attached to the host by means of a massive holdfast. It is the procambial initials which actually invade the host tissue. After penetrating the host, the individual procambium initials proliferate in the cortex, each giving rise to a cortical strand. It was concluded that the endophytic system of Arceuthobium cannot be interpreted in terms of classical concepts of plant homology; it is, in fact, an organ sui generis.     

EMBRYOGENY AND HISTOGENESIS IN NERIUM OLEANDER II. ORIGIN AND DEVELOPMENT OF THE NON-ARTICULATED LATICIFER

Mahlberg , P. G. (U. Pittsburgh, Pittsburgh, Pa.) Embryogeny and histogenesis in Nerium oleander. II. Origin and development of the non-articulated Iaticifer. Amer. Jour. Bot. 48(1): 90–99. Illus. 1961.—Laticifer initials, collectively considered as a laticifer system, are differentiated in the globular embryo from meristematic cells which occupy a position within the potential procambial tissue. A total of usually 28 initials, in Nerium oleander, arise as an irregular ring of cells directly below the embryonic shoot apex, during initiation of the cotyledonary primordia. No anastomoses occur between laticifer initials. During subsequent development of the embryo, the laticifer initials grow in a bi-directional manner and penetrate into the root, cotyledons and toward the shoot apex. Upon enlargement the initials bifurcate repeatedly, many branches penetrate into the cotyledons, others grow into the cortex of the hypocotyl or penetrate between cells of the procambium. Repeated nuclear divisions within each initial result in the formation of a multinucleated protoplast in this cell type. The tips of laticifers occupy intercellular spaces during their growth; they do not penetrate into or through adjacent cells. A plexus of laticifer branches is formed within the cotyledonary node of the mature embryo. No new initials are formed during subsequent growth of the plant, rather certain branches from the cotyledonary nodal plexus penetrate into the enlarging shoot system. The nature of their growth habit and branching suggests that the tips of laticifer initials exhibit an intrusive form of growth.     

SHOOT APEX ORGANIZATION AND ORIGIN OF THE RHIZOME-BORNE ROOTS AND THEIR ASSOCIATED GAPS IN DENNSTAEDTIA CICUTARIA

The shoot apex of Dennstaedtia cicutaria consists of three zones—a zone of surface initials, a zone of subsurface initials, and a cup-shaped zone that is subdivided into a peripheral region and central region. A diffuse primary thickening meristem, which is continuous with the peripheral region of the cup-shaped zone, gives rise to a broad cortex. The roots occurring on the rhizomes are initiated very near the shoot apex in the outer derivatives of the primary thickening meristem. The roots that occur on the leaf bases also differentiate from cortical cells. Eventually, those cortical cells situated between the newly formed root apical cell and the rhizome procambium (or leaf trace) differentiate into the procambium of the root trace, thus establishing procambial continuity with that of the rhizome or leaf trace. Parenchymatous root gaps are formed in the rhizome stele and leaf traces when a few of their procambial cells located directly above the juncture of the root trace procambium differentiate into parenchyma. As the rhizome procambium or leaf trace continues to elongate, the parenchyma cells of the gap randomly divide and enlarge, thus extending the gap.     

Late Embryogeny of Fokienia

The young embryo of Fokienia became massive and columnar in the middle of July in Chekiang, China. The polarity of the embryo has already been evident at this time. The cell in the free apex of the columnar embryo are smaller, while in the opposite end the cells continuous to the suspensor are larger in size and irregular in arrangement. Then a group of root initials appears at the middle part of the arc formed by the arrangement of the cells about 10 cells deep from the free apex. All kinds of tissues and organs of the embryo were differentiated in the first week of August. And the root initials become evident. Finally the root initials give rise to the procambium and the embryonic cortex upward and the root cap downward. There are about 20 layers of cells of the procambium in width and only about 10 layers of cells in cotyledonal procambium strand in the mature embryo. The cells of the embryonic cortex are continuous to those of the pericolumn of the root cap. The embryonic epidermis is absent in root cap. The embryo became fundamentally mature about the end of September. The hypocoty and cotyledons are well developed and each constitutes about 40% of the total length of the mature embryo. The root cap is rather weak, only about 10% of the total length. And the rest is the degenerated suspensors. The pith and secretory cells are absent in the mature embryo. The cotyledonal number of the embryo is 2. In mature ovule, there are 15 layers of nucellar cells in width in micropylar part but only 4—5 layers around the rest of the female gametophyte. The megaspore membrane is about 3.6μin thickness. When the young embryo is in the columnar stage, the nuclei of the female gametophytic cells are dividing and forming polynucleate cells. Thus, each cell usually has 2—4 nuclei. In this case, the cells of female gametophyte are large and isodiametric and about 60—120 μ in diameter. But the cells in the outer layer of the female gametophyte are rather small and they are usually uninuclear, rarely binuclear. The present article also deals with the starch distribution during the late embryogeny.     

THE DEVELOPMENTAL ANATOMY OF THE ANNONACEAE. I. EMBRYO AND EARLY SEEDLING STRUCTURE

Development of the epicotylary meristem is delayed; thus, it apparently has no immediate controlling influence on the procambium initiation either in the hypocotyl-root axis or in the cotyledons. Contrary to the situation found in most herbaceous dicotyledons, the procambium forms a hollow cylinder in the axis of the mature embryo. The first protophloem differentiates simultaneously in the upper part of the hypocotyl-root axis, at the base of the cotyledons, and in the cotyledons; protoxylem differentiation follows the same pattern as that of the protophloem. The direct vascular connection between the hypocotyl-root axis and cotyledons is established early in the mature embryo, and the fundamental aspects of transition phenomena are exhibited by the procambium before mature vascular elements differentiate. In the mature embryos, the pattern of lateral differentiation is irregular in that the number of protoxylem elements present at different levels is quite variable.     

LEAF HISTOGENESIS IN LACTUCA SATIVA WITH EMPHASIS UPON LATICIFER ONTOGENY

The leaf primordia of Lactuca sativa ‘Meikoningen’ develop from a subapical initial in the second layer of the tunica on the side of a fiat shoot apex. Subsequent growth of the subsurface lamina is initiated by submarginal initials which divide anticlinally to produce an adaxial layer and ***a biseriate abaxiallayer, and periclinally to produce a middle layer from which procambium differentiates. The protoderm is derived from the first tunica layer by continuous anticlinal divisions. The activity of the subapical and submarginal initials is completed when the leaf is 0.3 mm in length and 4.0 mm in width, respectively. Continued growth of the leaf to 130-150 mm results from intercalary cell division and enlargement. The mature venation is visibly delineated when the leaf is 25-30 mm in length. Laticifer and phloem cells are initiated by the same mother cells in the ***procambium. The former become non-septate laticifers by resorption of cross walls. They mature concurrently with the phloem and before the xylem.     

DEVELOPMENTAL ANATOMY IN ROOTS OF IPOMOEA PURPUREA. I. RADICLE AND PRIMARY ROOT

The developmental anatomy of the primary root of Ipomoea purpurea was studied at several growth stages, beginning with the radicle. The radicle is generally composed of three superimposed tiers of initials, which produce the vascular cylinder, cortex, and columella; and a peripheral band of lateral rootcap-epidermal initials. The radicular cortex contains 16–19 immature laticifers; none of the tissue regions in the radicle contains mature cells. Following germination and during the first 2–3 cm growth of the primary root the apical meristem and its derivative tissues undergo a series of modifications. Root apical diameter decreases as cells in lateral portions of the rootcap elongate; meanwhile, the columella enlarges vertically. The relationship between cortical and columellar initials changes as fewer mitoses occur in the former while the latter remain active. In longer roots the columellar initials are directly in contact with the vascular initials. Cortical size diminishes during early root growth as cortical laticifers and their associated cells cease to be produced by the outer cortical initials and ground meristem. Early procambium, at the level of vascular pattern initiation, decreases in diameter by cellular reorientation, and the vascular cylinder decreases in overall diameter although the tetrarch pattern remains unchanged.     

STUDIES ON THE SEED OF BANANA. I. ANATOMY OF THE SEED AND EMBRYO OF MUSA BALBISIANA

Mc Gahan , Merritt W. (United Fruit Co., Norwood, Mass.) Studies on the seed of banana. I. Anatomy of the seed and embryo of Musa balbisiana. Amer. Jour. Bot. 48(3): 230–238. Illus. 1961.—The seed coat of Musa balbisiana Colla consists of a relatively thick outer integument and a 2–cell-layered inner integument. The entire seed coat is sclerified, but routine tests for lignin are negative. Within the outer integument there is a zone of unusual sclereids tentatively termed “multiluminate.” Between the inner integument and the remnants of the nucellus is a cuticle 10–12 μ thick. The micropylar plug and collar are typical of the genus. The chalazal mass is an annular region of gelatinous cells. The mature embryo is comprised of a massive cotyledon, an epicotyl with 1 leaf primordium, a primary root primordium, and several adventitious root primordia. Procambium is well developed, but no mature vascular elements are present in the embryo.     

The ontogeny of the vascular cambium inGinkgo biloba roots

Observation was made on early ontogeny of vascular cambium in the developing root ofGinkgo biloba L. After completion of root elongation, the vascular meristem gradually acquires cambial characteristics. Strips of the periclinal division of cells in transverse section are observed on the inner side of phloem when the primary xylem and phloem in the stele have been established. The strips are united into a continuous layer between phloem and xylem. In tangenital section, the procambium shows a homogeneous structure, which is initially composed of short cells with transverse end walls and subsequently, of long cells with tapering ends. Then, the procambium is organized into two systems of cells; axial strands of short cells with transverse end walls resulting from the sporadic transverse divisions of long cells, and long cells with tapering ends. Still later, the short cells are divided frequently in a trasverse plane exhibiting one or a few cells in width and several decades of cells in height, while the long cells are elongated. The frequency of transverse divisions of the short cells decreases in subsequent stages. Eventually, the short cells in axial strands are vertically separated from one another by the elongation of neighboring long cells and by the decrease in the frequency of transverse divisions of short cells themselves. Cambial initials occur in two forms; ray initials a few cells in height and one cell in width derived from the short cells, and fusiform initials with tapering ends derived from the long cells.     

THE INITIATION OF SPOROPHYTES BY OBLIGATE APOGAMY IN CHEILANTHES CASTANEA

The initiation of apogamous sporophytes in Cheilanthes castanea was recorded by daily photography of individual gametophytes. Whereas an ordinary embryo arises from a zygote, apogamous embryos of C. castanea originate from one to three initial cells which occur just behind the apical region of the prothallus. The initial (or initials) produce cells with small chloroplasts behind the sinus of the gametophyte. The appearance of cells with smaller chloroplasts than those normally found in gametophytes is the first indication that apogamy is occurring. The cells with small plastids produce a group of densely-cytoplasmic meristematic cells. The size of the meristematic mass increases until shoot and root apices of the apogamous embryo are organized.     

THE ULTRASTRUCTURE OF THE QUIESCENT CENTER IN THE APEX OF CULTURED ROOTS OF CONVOLVULUS ARVENSIS L.

Cultured roots of the common bindweed, Convolvulus arvensis L. growing at the rate of 15–30 mm/day in sterile nutrient medium were fixed for electron microscopic analysis. The ultrastructure of the quiescent center, the initials of the ground meristem, and the initials of the procambium were studied in order to determine whether sequential structural changes could be correlated with models for specifying the mechanisms by which cell differentiation and cell division might be controlled. The differentiation of cells in the root proper occurs very gradually in linear files from the site of the quiescent center proximally into the different tissue regions. Major structural changes, such as the orientation and subsequent elongation of cells along the longitudinal axis of the root and cell wall changes, indicate that the control of differentiation and perhaps cell division occurs in radial gradients outwardly from the quiescent center.     

Early ontogeny of vascular cambium II.Aucuba japonica andWeigela coraeensis

InAucuba andWeigela the six vascular bundles distributed as a hexagon become connected tangentially by meristematic cells into a procambial cylinder in the early stage. In the tangential view, the procambial cylinder shows a rather homogeneous structure. InAucuba, some cells of the procambium elongate in a relatively earlier stage and the rest also elongate during subsequent stages. All of these cells have tapering end walls. Then some long cells divide transversely and form two systems in the vascular meristem, one made up of long cells and the other of short ones. The long cells become the fusiform initials and the short cells, the ray initials. InWeigela, the homogeneous procambium is organized in the later stages into two systems, one of long cells and the other of short cells in axial files. Most of the long cells have tapering end walls and the short cells transverse end walls. Some of the short cells elongate to intrude between adjacent cells and become long cells. The long cells become the fusiform initials. Radial divisions in some short cells occur occasionally. Some of these cells elongate and the rest remain in the axial files. Some short cells in the axial files are vertically separated from each other by the elongation of adjacent long cells. however, this occurs infrequently and the height of axial files is still several decades of cells. Short cells in axial files eventually become ray initials.     

Observations on phyllotaxis, stelar morphology, the shoot apex and gemmae of Lycopodium lucidulum Michaux (Lycopodiaceae)

Phyllotaxis in Lycopodium lucidulum consists of low alternating spirals, with the adult shoots corresponding to a system of 5 + 5 contact parastichies in which there are ten orthostichies. Each major stelar lobe is a sympodium of the leaf traces of two orthostichies and each lobe has two mesarch xylem poles, Differentiation of both the procambium and xylem of the leaf traces is bidirectional, that is differentiation first commences in the leaf base and then is acropetal into the leaf and basipetal into the stem. Furthermore, the procambium of the axis does not extend above that of the youngest leaf primordium and the axial procambium is in part a composite of that of the leaf traces. Thus, it is concluded that the stele in this taxon is not a strictly cauline structure. The shoot apex consists of four zones—a zone of surface initials, a zone of subsurface initials, a peripheral zone and a rib meristem. This zonation pattern is essentially the same as that of the seed plants. From their inception, gemma primordia also exhibit shoot apical zonation and are entirely different from leaves in their subsequent growth pattern and vascularization. Although the gemmae occupy leaf sites in the phyllotactic sequence, they are interpreted as arrested stem dichotomies on the basis of their development and vascular system.     

THE DEVELOPMENT OF THE SEED OF ABUTILON THEOPHRASTI I. OVULE AND EMBRYO

Winter , Dorothy M. (Iowa State U., Ames.) The development of the seed of Abutilon theophrasti. I. Ovule and embryo. Amer. Jour. Bot. 47(1): 8–14. Illus. 1960.—Abutilon theophrasti Medic, is a widespread annual weed which produces an abundance of seed in capsules which mature within 20 days after pollination. Ovule differentiation may be observed at least 8 days before anthesis when a sporogenous cell becomes evident and 2 integuments are initiated. An 8-nucleate embryo sac is produced from the chalazal megaspore approximately 2 days before anthesis. The outer integument of the mature campylotropous ovule consists of 2 cell layers, the inner integument has 6 to 15 cell layers. The initially free-nucleate endosperm becomes cellular betwen 3 and 7 days after pollination. At maturity a thin layer of gelatinous endosperm encases the embryo. The Asterad-type proembryo of Abutilon has a stout suspensor and develops rapidly. Four days after pollination cotyledons are initiated; 4 days later a leaf primordium is evident. Fifteen days after pollination the embryo, which has essentially completed its growth, consists of a large hypocotyl with root promeristem and root cap at its basal end, and 2 flat, folded, leaflike cotyledons enclosing a small epicotyl at its upper end. The epicotyl consists of an embryonic leaf and a stem apex.     

THE ROOT APEX OF MALVA SYLVESTRIS. I. STRUCTURAL DEVELOPMENT

The apical organization of the primary root of Malva sylvestris was analyzed at several growth stages, beginning with the embryo, to determine the structural changes that occurred during growth. Seeds were germinated, and plants were grown under controlled conditions. There were three discrete groups of initials in the embryonic root: those of the central cylinder, cortex, and secondary columella. The secondary columella initials consisted of a plate of cells flanked by a ring of cortical initials. The lateral portion of the rootcap shared a common origin with the epidermis. During growth both the initials of the secondary columella and outer cortex produced rootcap cells. The first indication of the outer cortical initials participating in rootcap formation was observed in roots 3 cm long. In 6-, 9-, and 16-cm roots the cellular continuity between the outer cortex and rootcap was marked, but in 23- and 33-cm roots the histogenic continuity between the outer cortex and rootcap was not evident. In all growth stages the initials of the central cylinder and inner cortex retained their histogenic integrity.     

THE DEVELOPMENTAL ANATOMY OF LONG-BRANCH TERMINAL BUDS OF PINUS BANKSIANA

A study of the composition of long-branch terminal buds (LBTB) of Pinus banksiana Lamb. and the yearly periodicity associated with their formation, development, and elongation was undertaken. Each LBTB has lateral bud zones and zones of cataphylls lacking axillary buds. When present, staminate cone primordia differentiate from the lowest lateral buds in the lowest lateral bud zone of the LBTB. Ovulate cone primordia and lateral long-branch buds can differentiate from the upper lateral buds in any lateral bud zone. When both types of buds are present, lateral long-branch buds are uppermost. Dwarf-branch buds occur in all lateral bud zones. During spring LBTB internodes elongate, new cataphylls are initiated, dwarf branches elongate, needles form and elongate, pollen forms and is released, and ovulate cones are pollinated. During summer buds form in the axils of the newly formed cataphylls. By early fall the new LBTB are in overwintering condition and the four types of lateral buds are discernable. The cytohistological zonation of the LBTB shoot apex is similar to that of more than 20 other conifer species. Cells in shoot apices of pine are usually arranged in distinct zones: apical initials, subapical initials, central meristem, and peripheral meristem. Periclinal divisions occur in the surface cells of the apex; therefore no tunica is present. At any given time, shoot apex volume and shape vary among LBTB in various positions on a tree. In any one LBTB on a tree, shoot apex shape changes from a low dome during spring to a high dome during summer to an intermediate shape through fall and winter.     

CYTOLOGY AND MORPHOGENESIS IN THE RHIZOMORPH OF ARMILLARIA MELLEA

Aided by the techniques of thin sectioning and electron microscopy, the apical region of the rhizomorph of Armillaria mellea has been examined. This region is composed of concentric zones of morphologically distinct tissues derived from a subapical meristematic zone designated the apical center. Meristematic activity is of two types: (1) primary, localized in the apical center, in which new hyphal elements are formed from apical initials, and (2) secondary, localized in the lateral regions of the apex, in which elaboration of the hyphal elements by means of elongation and secondary crosswall formation takes place. From these meristematic zones the tissues of the mature rhizomorph are derived and include: (a) peripheral hyphae, (b) cortex, (c) subcortex, and (d) primary and secondary medulla. The manner of differentiation of an apical initial appears unique and involves synchronous nuclear divisions accompanied by segmentation in many planes. The result of this activity is the formation of multinucleate hyphae. Apical initials are usually highly cytoplasmic and possess peculiar non-membrane-bound fibrous bundles, but in all other respects they resemble the hyphae of most Basidiomycetes thus far examined with the electron microscope.     

The Development and Structure of the Late Embryo in Torreya grandis

The materials used in this investigation were collected during 1980–1983 from Zhuji county of Zhejiang province, China. Seed of Torrcya grandis is an important dry “fruit” and used for edible oil. It is endemic to China. The primordia of male strobili are differentiated before October in the first year, while those of female strobili occur later. The microspore mother cells and megaspore mother cells are found in March and April in the second year respectively. The fertilization takes place in August and the dormant embryo overwinters at the proembryo stage. Eventually the proembryo begins to differentiate and its development starts in July of the third year. Thus the interval from fertilization to latembryogeny of Torreya grandis lasts for about 11 months. When the seeds of Torreya grandis are shed 'in August the embryo within the seed is still immature. It requires a period of after-ripening. The experiments show that the embryo resumes to develop and differentiate during 1–3 months in stratification in moist sands. The development and structure of late embryo are characterized as follows: 1. The cotyledon of the mature embryo in Torreya grandis is of 15000 μm in length and 87% of the embryo. The hypocotyl is vary shert and only 13% of the embryo. This kind of structure of the embryo in Torreya is very rare among conifers and in some degree similar to that of Keteleeria. When seed is shed the meristem of cotyledon is just differentiated and only 100–200 μm in length at the end of July to the middle of August. As the seeds are stratified in moist sands for 1–3 months, the cotyledon increases about 100 times than in room temperature in Zhuji county. 2. There is a large secretory canal in either side between the procambium and the cortex of the mature embryo. The secretory canal consists of epithelial cells of 4–5 layers. It is very peculiar in conifers. 3. The shoot apex does not begin to differentiate, until the seed has been fallen from the tree. 4. The column of the root cap is rather short and consists of the cells of about 10 layers in height and 6 layers in width. 5. Proteins are only found in the focal zone of the free apex of the young embryo but without any starch grains. The starch is abundantly distributed in the opposite end from the root initials down to root cap and the entire transitional zone. It is interesting to note that neither proteins nor starch grains are found in the suspensor system. It is assumed that the protein may be the main form of storing material in the actively growing cells and tissues of embryo in Torreya grandis.     

EMBRYO STRUCTURE AND STORAGE RESERVE HISTOCHEMISTRY IN THE PALM WASHINGTONIA FILIFERA

The seed of Washingtonia filifera (Lindl.) Wendl. is hemispherical and has a smooth testa. The embryo is located on the rounded side of the seed near the raphe. The embryo consists of a prominent single cotyledon, an epicotyl, and a small root apex. The shoot apex is oriented at a right angle to the long axis of the embryo and possesses 2 to 3 leaf primordia. The cotyledon functions as a storage organ and is composed of three cell types with similar ultrastructure. These three types—the parenchyma, protoderm, and procambium—can be distinguished on the basis of position, size, and shape. The procambial strands in the cotyledon consist of a ring of bundles grouped into two distinct sympodia and extend from beneath the shoot apical meristem to the tip of the cotyledon where they are situated very close to the surface. The most prominent organelles within all cell types are protein bodies, lipid bodies, and crystalline protein fibers. The protein bodies contain small crystalline inclusions which are presumed to be phytin. Protein bodies in the protoderm were smaller, denser-staining, and contained fewer crystalline inclusions than those in the parenchyma or procambium. On a volume basis, the parenchyma was shown to be 43% protein bodies, 25% lipid bodies, 15% cytoplasm, 7% cell wall, 4% intercellular space, 2% nuclei, and 4% other organelles (mitochondria and plastids).     

小麦种子根的发育解剖

小麦胚胎发育过程中通常形成5条幼根(少数可形成6条),这些根统称为种子根,中间最先发生的为初生根.初生根的原基在胚胎发育的早期就在胚轴的一侧发生,原基细胞由不规则到规则排列。侧生种子根的原基在胚胎发育后期才出现,通常成对发生,并且是由胚轴上的节(盾片节和胚芽鞘节)维管束外方的细胞形成。侧生种子根的发育明显较初生根的快,分化能力也较强,后生木质部导管母细胞出现早,数目较多.因此,小麦胚胎发育过程中从胚轴上形成的这些侧生的种子根,形态上,仍应看作是一些不定根,其结构特征与后来形成须根系的不定根的比较近似。