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.     

Morfogenesi in espianti caulinari di Aucuba japonica Thunb. coltivati in vitro

Abstract

Morphogenesis in caulinar fragments of Aucuba japonica Thunb. in vitro. — Caulinar fragments of Aucuba japonica Thunb. are grown in vitro in different media. They show an obvious polarity; in fact only the basal section surface shows a vigorous proliferation and is able to form roots. When this surface emerges it gives rise to a green luxuriant callus; when the same surface is immersed it forms a protuberance in the shape of crateriform invaginations. Our histological studies indicate not only the connection between new proliferations and the pre-existent vascular tissues, but also the areas where the pseudothalli originate. The changes in the cortex and the central cylinder which are caused by the in vitro cultures were observed.     

Occurrence of crystals in vascular cambium

Summary Crystals are of common occurrence in plant tissues and considered to be waste products resulting from active metabolism of plant cells. They mostly occur in ray and vertical parenchyma of wood and bark of angiosperm trees and often develop in tissues which soon cease to be functional. While studying the annual rhythm of cambial activity in two species of theVerbenaceae, Tectona grandis (Rao andDave 1981) andGmelina arborea (Dave andRao 1982), crystals of calcium oxalate have been observed in ray initials (Figs. 1–10). The crystals were identified as calcium oxalate after a convincing chemical behaviour towards dilute (10%) acetic, hydrochloric, and sulphuric acids. They appear colorless when viewed with brightfield illumination (Fig. 3), and show birefringence in polarized light (Figs. 1 and 6). Radial, tangential, and transverse sections of 10–15 m thick were used to study the crystals in polarized light with a Carl Zeiss Axiomat microscope.     

Shoot morphology of Aucuba japonica incurred by anisophylly: ecological implications

Anisophylly, having leaves different in size and/or shape, was quantified in adult Aucuba japonica and simulations were carried out to evaluate the effects of anisophylly on the extent of self-shading at the single-shoot level as well as at the whole-canopy level. Clear anisophylly was observed in the individual after switching from the single-stemmed juvenile stage to the multi-stemmed adult stage. In such plants, leaf area in the canopy abruptly increased. The effective display of adult foliage involved a variety of morphological changes in addition to anisophylly, most prominently reduction in leaf size compared to juveniles. The simulation results indicate that diversity of leaf size and shape is an effective means of minimizing self-shading as well as allowing the efficient exploitation of a larger canopy volume in adult plants. Anisophylly also increased the biomass use efficiency of individual plants at maturity. Taken together, having diverse leaf forms is superior to having a single leaf form for maximizing area acquisition and for efficiently filling the acquired area. We therefore conclude that the anisophylly expressed in A. japonica is adaptive.     

Effects of shading on photosynthesis characteristics of Photinia x frasery and Aucuba japonica var. variegata

This paper studied the effects of different shading (light transmittance 20%, 40%, 60%, and 100%) on the photosynthesis characteristics of two ornamental foliage plants Photinia x frasery and Aucuba japonica var. variegata. After shading for six weeks, the net photosynthesis rates of two plants measured ex situ under natural light enhanced, compared to those measured in situ, and, with the increase of shading degree, the net photosynthetic rates had an increasing trend, with the maximum being 9.7 micromol x m(-2) x s(-1) for Photinia x frasery and 8.3 micromol x m(-2) x s(-1) for Aucuba japonica var. variegata. In the meantime, the transpiration rates of the two plants increased significantly. Shading increased the chlorophyll a, b, and a+b contents and the chlorophyll/carotenoids ratio, decreased the chlorophyll a/b, but less affected the carotenoids content. The phenotypic plasticity index (PPI) of net photosynthesis rate and transpiration rate of Photinia x frasery and Aucuba japonica var. variegate was 2.08 and 3.21, and 0.55 and 1.60, respectively. The chlorophyll and carotenoids contents of the two plants were relatively stable, indicating the minor influence of external environment factors on pigments. Aucuba japonica var. variegata had a higher shading tolerance than Photinia x frasery.     

The vascular cambium: molecular control of cellular structure

Indeterminate growth and the production of new organs in plants require a constant supply of new cells. The majority of these cells are produced in mitotic regions called meristems. For primary or tip growth of the roots and shoots, the meristems are located in the apices. These apical meristems have been shown to function as developmentally regulated and environmentally responsive stem cell niches. The principle requirements to maintain a functioning meristem in a dynamic system are a balance of cell division and differentiation and the regulation of the planes of cell division and expansion. Woody plants also have secondary indeterminate mitotic regions towards the exterior of roots, stems and branches that produce the cells for continued growth in girth. The chief secondary meristem is the vascular cambium (VC). As its name implies, cells produced in the VC contribute to the growth in girth via the production of secondary vascular elements. Although we know a considerable amount about the cellular and molecular basis of the apical meristems, our knowledge of the cellular basis and molecular functioning of the VC has been rudimentary. This is now changing as a growing body of research shows that the primary and secondary meristems share some common fundamental regulatory mechanisms. In this review, we outline recent research that is leading to a better understanding of the molecular forces that shape the cellular structure and function of the VC.     

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.     

Plasticity in leaf-area density within the crown of Aucuba japonica growing under different light levels

We assessed leaf-area density (LAD; m² m⁻³) within the crown of Aucuba japonica (Cornaceae) growing under different light regimes and analyzed the components of crown architecture that most influenced variation in LAD. At a whole-crown level, extension-unit (EU) density (EUs/m³) had the greatest impact on LAD. The number of leaves per unit EU length and EU length had a wide range of impacts depending on the degree of crowding of foliage on the EU. Leaf size had a lesser impact on LAD. LAD was higher in the uppermost crown and declined towards the base. The non-uniformity of LAD among crown layers was much greater under high irradiance. Individuals under high irradiance achieved greater LAD by increased branching, well-marked EU dimorphism and a larger number of leaves per unit EU length; the reverse was true for the individuals under low irradiance. We identified two distinct modes of growth response to light regime. Under high irradiance, individuals responded by differential growth between the layers of crowns with the lower crown suppressed and growth in the upper crown increased. Conversely, shaded individuals did not respond by differential growth between crown layers.     

Seasonal changes in the ultrastructure of the vascular cambium of Robinia pseudoacacia

 The ultrastructure of the vascular cambium of Robinia pseudoacacia L. was examined in trunk tissues collected over a 2 ¹/₂ year period. During dormancy, fusiform cells are densely cytoplasmic with many small vacuoles and centrally located nuclei. Mitochondria are round to oval in sectional view. The plastids are variable in shape, have few internal membranes, and generally lack starch grains. The plasmalemma is smooth in outline. Proteinaceous material occurs in the vacuoles and many lipid droplets are scattered throughout the ground substance. Smooth tubular ER, often highly dilated, predominates, but short segments of rough ER are also present. Abundant free ribosomes are evenly distributed throughout the ground substance and the dictyosomes are inactive. Microtubules are parietal and have various orientations. During reactivation, the plasmalemma becomes irregular in outline and begins to form invaginations. Concurrently, the proteinaceous material disappears, the vacuoles begin to fuse, polysomes appear, and the dictyosomes begin to produce vesicles. During the period of cambial activity, fusiform cells are highly vacuolate, and the nuclei are centrally located. The mitochondria are round, oval, or elongate. Now the plastids contain phytoferritin, starch grains, or both. Many large invaginations of the plasmalemma intrude into the vacuole, pushing the tonoplast inward and pinching off into the vacuole, which lacks proteinaceous material. Lipid droplets are scarce. Most ER is rough, and ribosomes are generally aggregated as polysomes. Dictyosomes are actively producing vesicles. During the transition to dormancy, the fusiform cells gradually assume the appearance typical of the dormant cambium.     

A mathematical model of pattern formation in the vascular cambium of trees

The beautiful patterns apparent in wood grain have their origin in the alignment of fusiform initial cells in the vascular cambium of trees. We develop a mathematical model to describe the orientation of fusiform initial cells, and their interaction with the plant hormone indole-3-acetic acid (auxin). The model incorporates the following four assumptions: (1) auxin is actively transported parallel to the long axis of the initials, (2) auxin diffuses perpendicular to the long axis of the initials, (3) the initials tend to orient parallel to the flux of auxin through the cambium, and (4) adjacent initials tend to orient parallel to one another. Each assumption is justified on the basis of available evidence and cast in mathematical form. Our main result is a pair of nonlinear differential equations that describe the coupling between the distribution of auxin in the cambium and the orientation of fusiform initials. Numerical solutions to the equations show qualitative resemblance to the wood grain patterns observed at branch junctions, wounds and knots, and topological defects.     

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.     

Modelling for rearrangement of fusiform initials during radial growth of the vascular cambium in Pinus sylvestris L.

In contrast to common belief, recent studies have confirmed that intrusive growth of fusiform cambial initials has a significant role in the rearrangement of the initials, but does not contribute to the cambial circumference increment. We observed a rapid rearrangement of cambial initials on a long series of transverse sections of the vascular cambium and the wood of a 50-year-old pine (Pinus sylvestris L.) tree. A comparison of cell arrangement in consecutive sections, as well as a critical analysis of tangential reconstructions, has confirmed that changes in cell locations in a group of cells on the tangential surface caused no change in the total tangential width of the whole group. Models illustrating changes in locations of the initials have been proposed, assuming that intrusive growth, which makes the growing initials intrude between the neighbouring initials and their immediate derivatives, is localized on the longitudinal edges of cells. We infer that intrusive growth of the cambial initials in P. sylvestris is not involved in the cambial circumference increment, but plays a significant role in the rearrangement of the initials, probably allowing for a relaxation of shearing strains generated during radial growth. The relationship of intrusive growth with the elimination of initials has been discussed with reference to the frequency of anticlinal divisions. It has been proposed that the occurrence of anticlinal divisions in excess over the actual requirement for increase in the cambial circumference could be due to internal shearing strains.     

Early larval ontogeny of the Permo-Carboniferous temnospondyl Sclerocephalus

The early larval development of the temnospondyl Sclerocephalus sp. is analyzed, based on 38 specimens from the Lower Rotliegend (Permo-Carboniferous boundary) of the Saar-Nahe Basin (south-west Germany). The study focuses on the smallest larval specimens, which exemplify changes in both proportions and ossification patterns. In comparison with dissorophoid larvae, the skull ossifies more fully and at a much faster rate; the smallest specimens already have completely formed circumorbital bones that are sutured throughout. Sculpturing undergoes two marked changes, first from uniformly pitted to pits of variable size and regional differentiation, and finally to the origin of ridges. The palate of small larvae differs from that of larger specimens in patterns of dentition, having more teeth including a denticle field on the cultriform process. The mandible of small larvae is described for the first time, being narrower than in adults and having three dentigerous coronoid elements. The smallest specimens have poorly ossified neural arches, lack vertebral centra, and have faintly ossified humeri, femora, and very poorly developed distal elements. The posterior ribs, metapodia, and phalanges appeared after the dermal elements of the pectoral girdle, whereas the scapulocoracoid and ischium are absent throughout the larval period. Early growth and differentiation of the limbs and the ilium illustrates the developmental patterning of the appendages, which proceeded from proximal to distal. Dermal squamation is uniform in small stages, consisting of round or oval osteoderms with pronounced growth rings; in large larvae, they start to differentiate in certain body regions.     

Estimation of directional division frequencies in vascular cambium and in marginal meristematic cells of plants

Using simple arithmetical formulae, it is shown that, when the meristematic initial cells of a growing plant organ are arranged in a ring, the cellular dimensions predict the relative frequencies of anticlinal and periclinal divisions which these cells undergo. The pattern of cell file branching which appears during the course of development, and which is predicted by this mathematical model, is validated using data pertaining to the numbers and dimensions of initial cells within the secondary vascular cambium of hybrid aspen trees. Data pertaining to a second, simpler set of initial cells which comprises the outer cellular ring of the thallus of the alga Coleochaete orbicularis, and from which all the radial cell files of the circular disc-like thallus are descended, have also been used for model validation. Combining the mathematical approach to division frequencies with data of actual cell sizes permits inferences about the course of the increase of the number of cell files (generated by the anticlinal divisions) and the number of cells within each file (generated by the periclinal divisions) during the earlier stages of secondary tissue or thallus development, and also about how they will develop at future stages. The question whether or not cell division patterns conform to the geometry of the system in which the cells are embedded is also discussed.     

Early ontogeny of the human femoral bicondylar angle

The presence of a femoral bicondylar angle consistently and significantly greater than 0° has been a hallmark of hominid bipedality, but its pattern of development has not been documented. We have therefore compiled cross-sectional data on the development of the articular bicondylar angle for a clinical sample of modern humans and of the metaphyseal bicondylar angle for two Recent human skeletal samples, one predominantly European in origin and the other Amerindian. All three samples exhibit a pattern of a bicondylar angle of 0° at birth and then a steady average increase in the angle from late in the first year postnatal, through infancy, and into the juvenile years. The two skeletal samples reach low adult values by approximately 4 years postnatal, whereas the clinical sample with a lowered activity level appears to attain consistent adult values slightly later (approximately 6 years postnatal). In addition, two modern human individuals, one nonambulatory and the other minimally ambulatory, show no and little development, respectively, of a bicondylar angle. These data, in conjunction with clinical and experimental observations on the potential and form of angular changes during epiphyseal growth, establish a high degree of potential for plasticity in the development of the human bicondylar angle and the direct association of a bipedal locomotion and (especially) posture with the developmental emergence of a human femoral bicondylar angle. © 1994 Wiley-Liss, Inc.     

Comparative anatomy and ultrastructure of active and dormant vascular cambium in rhizome ofBotrychium ternatum

Vascular cambium ofBotrychium ternatum rhizome varied according to age, position and season was studied by light and electron microscopy. Cambium at the 6th internode (6-year-old cambium) had the greatest number of active cambial cells in August and September, thus it was in the most active stage. The active cells were characterized by the presence of a large vacuole, few storage materials such as starch grains within plastids or lipid droplets, a thin tangential wall; and various cell organelles in the thin peripheral layer of cytoplasm. When the 6-year-old cambium reached its dormant season after November, the dormant cells were filled with numerous storage materials and had few cell organelles. Our observations suggested that the initiation and cessation of cambial activity may be correlated with the annual life cycle of this plant: the vegetative and reproductive leaves began to emerge in June and July, respectively, and the sporophyll withered in November after the spore dispersal. Most cambial cells at the 10th internode, which remained in a dormant state throughout the year, were filled with numerous storage materials. Our results indicated that the activity of vascular cambium in the 10th internode was determinate.     

Early ontogeny of monocytes and macrophages in the pig

Prenatal development of cord blood monocytes and tissue macrophages was studied in pig foetuses by immunophenotyping and functional assays. The function of peripheral blood monocytes was compared in germ-free and conventional piglets. First macrophages were identified by electron microscopy in foetal liver on the 25th day of gestation. Monoclonal antibodies against porcine CD45 and SWC3 antigens were used for flow cytometric identification of myelomonocytic cells in cell suspensions prepared from the yolk sac, foetal liver, spleen and cord blood. Leukocytes expressing the common myelomonocytic antigen SWC3 were found in all organs studied since the earliest stages of development. Opsonized zymosan ingestion assay was used to determine the phagocytic capacity of foetal mononuclear phagocytes isolated from cord blood, liver and spleen. In the foetal liver, avid phagocytosis of apoptic cells had been found to occur before cells were able to ingest zymosan in vitro. The first cells capable of ingesting zymosan particles were found on the 40th day of gestation in umbilical blood and 17 days later in foetal spleen and liver. Their relative proportion increased with age. Cord blood monocytes and peripheral blood monocytes in germ-free piglets had low oxidatory burst activity as shown by iodonitrophenyl tetrazolium reduction assay. A remarkable increase of oxidatory burst activity was observed in conventional piglets, probably due to activation of immune mechanisms by the microflora colonizing gastrointestinal tract.