Early ontogeny of vascular cambium III

Observations were made on structural changes from the procambium to cambium in the developing shoots ofRobinia pseudo-acacia andSyringa oblata, both of which are characterized by relatively short fusiform initials. In both species, the procambium in transverse view shows radial seriations of cells as a result of repeated tangential divisions, and there is an almost continuous procambial cylinder in the young stem in the earlier stage. The procambium in tangential view has initially a homogeneous structure and later develops into two systems, one made of long cells, the other of short cells. Some of the short cells elongate to intrude among neighbouring cells and some long cells divide radially as well as tangentially. InRobinia, long cells have transverse or tapering end walls at a relatively earlier stage and mainly tapering end walls in the subsequent stages. Although some of the short cells develop into long cells, the height of axial files of the short cells decreases only a little, because of subsequent transverse divisions and growth of cells. InSyringa, long cells have mainly transverse end walls at a relatively earlier stage and usually tapering end walls in the subsequent stages. Short cells in axial files have predominantly transverse end walls. A number of additional long cells are derived from elongating short cells in the later stages. Therefore, the height of axial files becomes apparently lower than that of earlier stages. Radial divisions in short cells occur to some extent. Results are discussed in relation to the structure of the vascular meristem inGinkgo, Aucuba, Weigela, and others.     

喜树原形成层到形成层转化的研究

观察了枣树茎中原形成层到形成层的转化过程。距茎端0．5mm处,节间开始伸长之前,横切面上4—5个原形成束及束间的分生组织组成原形成层环。径向切面观,原形成层环呈现出较均一的结构。随着节间开始伸长,由于原形成层细胞发生假横向分裂,出现了长短两类细胞,长细胞多数端壁倾斜,短细胞多数端壁平截。以后,长细胞发育为纺缍状原始细胞,短细胞发育为射线原始细胞,部分射线原始细胞可以伸长井侵入生长而转化为纺缍状原始细胞。在节间伸长将停止时,此种转化基本完成。喜树为非叠生形成层,纺缍状原始细胞和射线原始细胞都有多核现象发生。     

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.     

Early ontogeny of vascular cambium

Observations were made on structural changes of procambium and the subsequent appearance of cambium in the developing shoot. The procambium in early stages shows radial seriation of cells as a result of repeated tangential divisions. In tangential view the procambium has initially a rather homogeneous structure and later becomes organized into two distinct systems, one composed of long cells and the other of short cells. The latter cells are arranged tangentially in axial files and transversely in radial files. They show repeated transverse divisions. Some of the short cells elongate to intrude between neighboring cells. Therefore, long cells may be derived both from cells of homogeneous structure in the first stage and from elongation of some of the short cells in axial files. Long cells have mostly tapering end walls and elongate actively. In the subsequent stages, the frequency of transverse divisions of short cells in axial files decreases and these cells expand radially. Short cells in axial files are separated from one another vertically by the elongation of neighboring long cells which break up the axial files as seen in tangential view. The vascular meristem in this stage is believed to initiate the cambium. Eventually, short cells remain mostly single, or form files of two or three cells in height in tangential view. The observations are discussed in relation to the structure of the vascular meristem in other plants.     

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.     

STRUCTURE AND ONTOGENY OF LATICIFERS IN CICHORIUM INTYBUS (COMPOSITAE)

The branched anastomosed laticifer system in the primary body of Cichorium intybus L. originates in embryos from files of laticiferous members at the boundary between phloic procambium and ground meristem. Upon seed germination, laticiferous members develop perforations in the end walls which become entirely resorbed. Perforations also develop in the longitudinal walls of contiguous laticiferous members and from lateral connections between developing laticifer branches. Additional laticiferous members originate as procambium differentiation proceeds, and their differentiation follows a continuous acropetal sequence in leaf primordia of the plumule. In roots, laticifers closely associated with sieve tubes in the secondary phloem originate from derivatives of fusiform initials in the vascular cambium. These laticifers develop wall perforations and in a mature condition resemble laticifers in the primary body. As the girth of the root increases, laticifers toward the periphery, unlike associated sieve tubes, resist crushing and obliteration. Laticifers vary in width from about 4 to 22 μm; the widest ones occur in involucral bracts and the narrowest ones in florets. There was no evidence that intrusive growth occurs during development of the laticifer system, although such growth may occur during development of occasional branches which extend through ground tissue independent of phloem and terminate in contact with the epidermis. Presence of amorphous callose deposits is related to aging of laticifers and mechanical injury.     

Rearrangement of cells in storeyed cambium of<Emphasis Type="Italic"> Lonchocarpus sericeus</Emphasis> (Poir.) DC connected with formation of interlocked grain in the xylem

The history of cellular events in the storeyed cambium of Lonchocarpus sericeus (Poir.) DC was analysed on the basis of changes in the cell arrangement in successive layers and strata of axial parenchyma in the xylem. The mechanism of formation of the regular interlocked grain was investigated. Inclination of fusiform cells changes intensively whereas height and position of storeys in the successive layers of axial parenchyma are constant. As a result, new contacts between cells are formed by means of the intrusive growth of ends of cells belonging to one storey between the tangential walls of cells of the neighbouring storey and unequal periclinal divisions, which give a new shape to the initials. The concept of intrusive growth between the radial walls of the fusiform initials in the formation of xylem with interlocked grain should be revised on this basis.     

Mathematical Modeling of Intrusive Growth of Fusiform Initials in Relation to Radial Growth and Expanding Cambial Circumference in Pinus sylvestris L.

This study on the cambium of Pinus sylvestris L. examines the intrusive growth of fusiform cambial initials and its possible contribution to the tangential and radial expansions of the cambial cylinder. The location and extent of intrusive growth of the fusiform initials were determined by microscopic observations and by mathematical modeling. In order to meet the required circumferential expansion of the cambial cylinder, the fusiform initials grow in groups by means of a symplastic rather than intrusive growth, leaving no room for the assumption that intrusive growth of the initials takes place between radial walls and has a direct role in the increase of the cambial circumference. Therefore, it is postulated that the fusiform initials grow intrusively between the tangential walls of the neighboring initials and their immediate derivatives and not between the radial walls of the adjacent initials as per common belief.     

The origin and development of vascular cambium in girdled stems ofEucommia ulmoides Oliv.

We conducted anatomical studies of girdled stems ofEucommia ulmoides at various developmental stages to elucidate the origin and development of callus and the vascular cambium. In the transverse view, ray initial cells in the cambial zone began to divide both periclinally and anticlinally 2 d after girdling. Fusiform initial cells started to enlarge at 3 d, then gradually proliferated via periclinal divisions. Thus, the callus was derived from the ray initial cells of the cambial zone as well as from fusiform initial cells. In the tangential view, callus cells derived from ray initial cells were short while those from fusiform initial cells were long, thereby producing a heterogeneous structure. However, the fusiform initial cells underwent transverse divisions 10 d after girdling, which resulted in shorter cells and a homogeneous callus structure. Afterward, some short cells divided transversely while others elongated, so that a heterogeneous form was regained. Finally, the vascular meristem that was girdled early in its development redifferentiated from short and long cells in the callus. The long cells developed into fusiform initials, with the short ones becoming ray initials.     

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.     

Early Development and Ultrastructure of the Major Veins and Their Sheath Tissues in the Maize Leaves

The report described the ultrastructural changes that occurred in the major veins and their associated bundle sheaths (BS) of the maize ( Zea mays L. ) leaf blade in the process of their differentiation from three adjacent cells in the middle layer of the ground meristem, the minimal number of cells involved with the initiation of a procambial strand and the associated BS. The inner cell underwent two successive unequal periclinal divisions: a smaller cell that later differentiated into the adaxial BS cell precursor, and a larger one that divided once again periclinally yielding an abaxial BS cell precursor and a centrally located procambial initial cell. One of the two lateral cells immediately adjacent to either side of the inner cell also divided periclinally; these derivatives, along with another lateral cell of the original three-celled unit formed the precursor cells of the lateral BS. Prior to the initiation of protophlcem differentiation, all of the procambial cells showed ultrastructural characteristics basically similar to the procambial initial. They possessed a prominent nucleus with electron-dense aggregates of heterochromatin, a dense cytoplasm rich in ribosomes, proplastids and mitochondria; also a thin wall containing numerous plasmodesmata. In many cases, only short pieces of rough endoplasmic reticulum cistemae and a few small sized vacuoles were present. In adclifton, evidence of cytoplasmic disintegration leading to new vacuole formation was noted in the process of proeambium development. It was observed that certain endoplasmic reticulum was engaged in the sequestration and lysis of cytoplasm. No apparent uhrastmctuml difference was found between the BS cell precursors and the procambial initials, that was, the distinction between the procambium and the surrounding BS cells occurred gradually after vein initiation, The major ultrastmctural changes which occurred during the differentiation of the meristematic BS cells into the vacuolated cells were (1) a proplastid to chloroplast transformation going through a prolamellar body stage, and (2) the appearance of the multi-concentric membrane complex which might play a role in the degradation of some ribosomes and other cytoplasmic components during the differentiation of BS cells.     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

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.     

Anatomy of the vascular cambium of Acacia nilotica (L.) Del. var. telia Troup (Mimosaceae) in relation to age and season

the majority of fusiform initials are multinucleate, a few having as many as eight nuclei. Their length increases down the stem from the apex, attaining a maximum in the old trunk and declining slightly near the base. The width of the initials exhibits similar variation. In the main trunk, fusiform initials, relatively short at the time of cambial reactivation (April), elongate steadily until July. There is a sharp decline in August/September, the cell length recovering during the winter. Seasonal variation in cell width is inconsistent. Ray cell initials, on the other hand, do not vary much in size. They divide more frequently in the older stem, adding to the size of rays. In young shoots, short and uni- to biseriate rays are most abundant, whereas tall and multiseriate rays dominate the cambial surface in the trunk region throughout the year, with their minimum population in the early phase of cambial activity and the maximum during peak activity. The overall proportion of fusiform initials in the cambial cylinder initially increases with age, from young shoots towards the base, and later becomes more or less constant in the trunk region. Here it remains noticeably high during the active growth period and relatively low for the rest of the year.     

Anatomy of the vascular cambium of Acacia nilotica (L.) Del. var. telia Troup (Mimosaceae) in relation to age and season

the majority of fusiform initials are multinucleate, a few having as many as eight nuclei. Their length increases down the stem from the apex, attaining a maximum in the old trunk and declining slightly near the base. The width of the initials exhibits similar variation. In the main trunk, fusiform initials, relatively short at the time of cambial reactivation (April), elongate steadily until July. There is a sharp decline in August/September, the cell length recovering during the winter. Seasonal variation in cell width is inconsistent. Ray cell initials, on the other hand, do not vary much in size. They divide more frequently in the older stem, adding to the size of rays. In young shoots, short and uni- to biseriate rays are most abundant, whereas tall and multiseriate rays dominate the cambial surface in the trunk region throughout the year, with their minimum population in the early phase of cambial activity and the maximum during peak activity. The overall proportion of fusiform initials in the cambial cylinder initially increases with age, from young shoots towards the base, and later becomes more or less constant in the trunk region. Here it remains noticeably high during the active growth period and relatively low for the rest of the year.     

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.     

Organisation of vascular cambium during different seasons in some tropical timber trees

The vascular cambium of Albizzia, Tectona, Terminalia, Calophyllum, Mangifera and Morinda was non-storied while that of Dalbergia was semi-storied. The ray initials were uniseriate in Terminalia and Calophyllum , both uniseriate and biseriate in Albizzia, Dalbergia, Mangifera and Morinda and were also multiseriate in Tectona . They were homogeneous in Albizzia and heterogeneous in the other species. The radial walls of cambial cells were always beaded, although beads were more prominent and closer to one another during periods of cambial dormancy than during activity. The fusiform initials were comparatively less vacuolated during dormancy/ least activity. When active, the fusiform and ray initials of all species, except Calophyllum , also showed multinucleate condition (2–10 depending on species). The proportion of ray initials to fusiform initials was almost constant throughout the year in all species.     

DEVELOPMENTAL CHANGES IN THE VASCULAR CAMBIUM OF POLYGONUM LAPATHIFOLIUM

Developmental changes in the vascular cambium of Polygonum lapathifolium were determined primarily by an analysis of the secondary xylem. The cambium and xylem consist of fascicular and interfascicular regions in this herbaceous dicotyledon. Near the pith vessels are restricted to the fascicular regions of the xylem. During secondary growth vessels are formed in some radial files in the interfascicular regions. Anticlinal divisions are of two types, oblique and lateral. In interfascicular files consisting of fibers only, about two-thirds of the anticlinal divisions are oblique. The oblique partition averages 31% of the length of the dividing initials. In interfascicular files consisting of vessel elements and fibers, there are almost equal numbers of oblique and lateral divisions. The oblique partition averages 37% of the length of the dividing initials in these files. Lateral divisions account for approximately three-fifths of the anticlinal divisions in the fascicular regions, consisting of vessel elements and fibers. The partitions formed in oblique anticlinal divisions average 64% of the length of the dividing cells in the fascicular regions. The frequency of anticlinal division is much higher in files consisting of vessel elements and fibers than in those consisting of fibers only. There is no loss of fusiform initials, except by ray formation. Ray initiation occurs by simple subdivision of fusiform initials. The findings are discussed in relation to the developmental changes in the vascular cambium in plants of different habits.