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.     

Structure of the vascular cambium of varying age and its derivative tissues in the stem of Ficus rumphii Blume

AJMAL, S. & IQBAL, M., 1992. Structure of the vascular cambium of varying age and its derivative tissues in the stem of Ficus rumphii Blume. The cambial cells of Ficus rumphii and their derivatives vary in size and number with the age of the tree. The fusiform initials, vessel elements, sieve-tube members, xylem fibres and phloem fibres are relatively short in the terminal part of the stem axis, consistently longer down the stem reaching a maximum, and short again in the main trunk. The width of the respective cells shows a similar variation. The ray cell initials increase both in number and size, and form rays of varying dimensions. Uniseriate and biseriate cambial rays are abundant, while multiseriate rays are especially sparse in the young shoot. The proportion of uni- and biseriate rays falls in the lower part of the stem whereas that of multiseriate rays increases. Likewise, short cambial rays are abundant in young branches but their frequency decreases in the trunk region, leading to a rise of the tall ray population. The rays in the phloem and xylem regions reflect a similar variation pattern with certain fluctuations. The overall proportion of the rays increases in the older part of the axis, reaching a maximum tangential area in the trunk. The transectional area of vessel elements remains more or less constant down to the base after an initial increase in the terminal region, while the sieve-tube proportion tends to be constant in the basal region after a consistent gain from the tree top basewards.     

Within-tree variation in cambial anatomy and xylem cell differentiation in Eucalyptus globulus

Summary Microscopic examination of the vascular cambium of Eucalyptus globulus, from four trees, at nine height levels, revealed a non-storied cambial structure with both ray and fusiform initials. Average fusiform initial length increased from 443 m at 70% of tree height from the base to a maximum of 484 m at 10% of tree height. At 2.5% of tree height average fusiform initial length sharply declined to 438 m. A strong positive correlation (r = 0.757, df = 7, P <0.05) existed between average fusiform initial length and average fibre length at each height, giving quantitative support to the notion that variation in fusiform initial length is an important mechanism influencing fibre length in hardwoods. However, 62% of the longitudinal variation in average fibre length was explained by the extent of elongation during differentiation, which increased linearly down the stem. At 70% of tree height, the average length of fibres was 1.84 times that of the fusiform initials, compared to 2.11 times at 2.5% of tree height. Concomitant with this longitudinal gradient of elongation was a gradient of increasing tangential area of the cambium occupied by cambial rays. This was accounted for by dimensional increases in the ray initial cells with decreasing height. Possible mechanisms that regulate the elongation of developing fibres and the proportion of cambial rays are discussed.     

Influence of exogenous ethylene on cambial activity,xylogenesis and ray initiation in young shoots of Leucaena leucocephala (lam.) de Wit

The effect of exogenous ethephon on cambial activity, xylem production and ray population in young shoots of Leucaena leucocephala was investigated anatomically. The application of ethephon showed a diphasic effect on cambial activity and xylogenesis in a dose dependent manner. Lower concentration of ethephon enhanced cambial activity while high concentrations reduced cambial cell divisions and daughter-cell differentiation. High ethephon concentration also resulted in shorter vessel elements, thick walled fibers and phenolic accumulation in ray cells and vessel elements, whereas low concentration allowed elongation of fibers and vessel elements. The density of rays increased significantly with increase in ethylene concentration. The evaluation of longitudinal sections of cambial zone in ethephon treated plants showed high frequency of transformation of fusiform initials into ray initials through divisions and segmentation, resulting in high ray frequency in both xylem and phloem. The present study demonstrates that ethylene plays an important role in regulating secondary vascular tissue composition by reducing the population of fusiform initials in the cambium.     

Cambial activity,annual rhythm of xylem production in relation to phenology and climatic factors and lignification pattern during xylogenesis in drum-stick tree (Moringa oleifera)

The interrelationship among seasonality of cambium, wood formation, cell size variation, lignification, tree phenology and climatic factors has been examined in Moringa oleifera, a tropical evergreen tree. The vascular cambium in Moringa is a storied with a distinct seasonal variation in its structure due to dimensional changes in rays. Though cambium remains active throughout the year it is sensitive to water availability. Peak cambial cell division and rate of xylem differentiation are influenced by average rainfall during the monsoon period. Cambial cell division reaches higher up in the tree trunk when it is supporting a high number of branches and leaves. Statistical analysis of cell size variation and climate factors revealed that xylem cell development is greatly influenced by rainfall and rarely by temperature. Lengths of fusiform initials and vessel elements are positively correlated. The pattern of lignification during xylogenesis shows that the vessels are the first element to develop lignified walls and ray cells are the last elements to become lignified. Fiber cell walls show more syringyl lignin, while the cell walls of other xylem elements are characterized by relatively more guaiacyl lignin units.     

Development of cambium and length of vessel elements and fibers in dwarfAlnus hirsuta (Spach) rupr

In a comparison of cambial cells and their derivatives between naturally occurring dwarf and normal trees, vessel elements and fibers in the annual rings of dwarf trees were found to be shorter, narrower and fewer than those of normal trees. The frequency of anticlinal divisions and loss of cambial initials were low during the differentiation of xylem cells from cambial initials in dwarf trees. The length and intrusive growth of fusiform initials were slightly less than those of normal trees. Thus, it was concluded that the shortening of vessel elements and fibers in dwarf trees was due to the fact that cambial initials were themselves shortened and underwent inactive intrusive growth during differentiation of the xylem mother cells.     

Seasonal histochemical changes in the cambium ofTectona grandis L. f. andgmelina arborea roxb

Starch, lipids and proteins are localized histochemically in fusiform and ray initials ofTectona grandis andGmelina arborea during the periodicity of cambium in the year 1978. The histochemical variations are correlated with the seasonal activity and dormancy of the cambium and phenology. Starch grains appear in the cambial cells with the development of young leaves after defoliation. Lipid and protein bodies are significantly high in dormant cambial cells and scarce in active ones.     

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.     

Seasonal Rhythms of Structure and Behaviour of Vascular Cambium in Ficus rumphii

Cambial structure and activity of Ficus rumphii Blume vary withthe changes in local climate. The cambial cells start swellingearly in April prior to the onset of periclinal divisions whichare most frequent in August. Cell division stops in October.During the growth season, initiation as well as cessation ofthe phloem production precedes that of xylem. A moderately hightemperature is correlated with the cambial reactivation. Onceinitiated, the activity continues at relatively low temperatures.Hot and dry environment favours the phloem production, whereashot and moderately humid conditions induce xylogenesis. Thesize and relative proportion of cambial initials also changewith season. Fusiform initials are shorter and broader duringthe rainy season (July–September) than for the rest ofthe year. Multiseriate and triseriate rays, as also the tallrays, outnumber the other types of rays throughout the year. Ficus rumphii, vascular cambium, phenology, climatic variation     

Developmental and Dimensional Changes in the Cambial Fusiform Initials ofHoloptelea integrifolia (Roxb.) Planch.

Divisional activity, intrusive growth of the cell wall and loss of fusiform initials have been studied in Holoptelea integrifolia. The dimensional changes in relation to mean length, length frequency, mean width and length variation in relation to fibre length have also been analysed.     

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.     

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.     

Developmental anatomy of vascular cambium and periderm ofBotrypus virginianus and its bearing on the systematic position of ophioglossaceae

The developmental anatomy of the vascular cambium and periderm ofBotrypus virginianus was studied, and its bearing on the systematic position of Ophioglossacease is discussed. The cambial zone including cambium is initiated in a procambial ring of the stem before primary vascular tissue is well differentiated. The presumed cambium is composed of fusiform and ray initials. The cambium is extremely unequally bifacial, producing secondary xylem centripetally, and quite a small number of parenchymatous cells but no secondary phloem centrifugally. The cambial activity persists long, although it is very low in the mature part of the stem. It seems that the circumferential increase of the cambium is accommodated by an increase in the number of cambial initials. Secondary xylem is nonstoried and composed of tracheids with circular-bordered pits with evenly thick pit membranes, and uniseriate or partly biseriate radial rays. It makes up the bulk of the stem xylem. Periderm is formed almost entirely around the stem, simultaneous with its increment due to the secondary xylem. The combination of these anatomical features of secondary tissue supports the idea that Ophioglossaceae are living progymnosperms.     

VASCULAR CAMBIUM AND WOOD DEVELOPMENT IN CARBONIFEROUS PLANTS. I. LEPIDODENDRALES

Patterns of activity in the vascular cambium of Carboniferous arborescent lycopods (Paralycopodites and Stigmaria) were studied by analysis of serial tangential sections of the secondary xylem. The analysis assumes that cell patterns in the wood accurately reflect those of the corresponding cambium. An evaluation using indirect evidence indicates that the assumption is valid as far as can be determined from comparison with living plants. The tracheids of the secondary xylem enlarge in a centrifugal pattern, suggesting a progressive enlargement of the fusiform initials. There is no evidence of periodic anticlinal division of these initials, and it is proposed that the increase in cambial circumference was accommodated primarily by an increase in fusiform initial size. In some axes with abundant secondary xylem there is evidence that isolated initials or groups of initials sporadically subdivided to form numerous, spindle-shaped meristematic cells. Some of these cells subsequently developed into typical cambial initials. Tissues presumably formed during the cessation of cambial growth in Lepidodendron and Stigmaria are described; the structure of the tissues is suggestive of a postmeristematic parenchymatous sheath. It is concluded that cambial activity in these arborescent cryptogams was clearly different from that of modern seed plants, further attesting to the distinctive nature of this ancient group.     

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.     

The anatomy of the chi-chi of <Emphasis Type="Italic">Ginkgo biloba</Emphasis> suggests a mode of elongation growth that is an alternative to growth driven by an apical meristem

The chi-chi of Ginkgo biloba L. are cylindrical woody structures that grow downwards from the branches and trunks of old trees, eventually entering the soil where they give rise to adventitious shoots and roots. Examination of segments of young chi-chi taken from a mature ginkgo tree revealed an internal woody portion with irregular growth rings of tracheid-containing secondary xylem covered by a vascular cambium and bark. The cambium was composed of both fusiform cells and parenchymatous ray cells. Near the tip of the chi-chi, these two types of cambial cells had orientations ranging between axial, radial and circumferential with respect to the cylindrical form of the chi-chi. The xylem rays and tracheids that derived from the cambium showed correspondingly variable orientations. Towards the base of the chi-chi, the fusiform cells and young tracheids were aligned parallel to the axis, indicating that the orientation of the cambial cells in basal regions of the chi-chi gradually became normalised as the tip of the chi-chi extended forwards. Nevertheless, in such basal sites, tracheids near the centre of the chi-chi showed variable orientations in accordance with their mode of formation during the early stages of chi-chi development. The initiation of a chi-chi is proposed to derive from a localised hyperactivity of vascular cambial-cell production in the supporting stem. The chi-chi elongates by tip growth, but it does so in a manner different from organ growth driven by an apical meristem. It is suggested that the chi-chi of Ginkgo is an “evolutionary experiment” that makes use of the vascular cambium, not only for its widening growth but also for its elongation.     

Seasonal variations in the cambial anatomy of Tectona grandis (Verbenaceae)

A study of seasonal activity of the cambium in Tectona grandis L. f. has shown that the activity initiates in the first week of June, reaches a peak in July and then slowly declines. The length of fusiform cambial initials undergo considerable variations during the activity and dormancy of the cambium. The initiation of cambial activity is closely associated with the opening of the dormant foliar buds in the first week of May. Cambium is more active with high numbers of immature xylem and phloem elements from July to September when the trees are with mature foliage and flowers and dormant from January to April when leaves dry and defoliation takes place. The differentiation of xylem and phloem starts simultaneously and the number of their immature elements reach the maximum in July.     

Variations in the Lengths of Fusiform Cambial Cells and Vessel Elements in Kalopanax pictus

Samples of a mature specimen of Kalopanax pictus, a ring-poroushardwood, were studied to compare the respective lengths offusiform cambial cells and vessel elements in the stem. Thelengths of dormant and reactivated fusiform cambial cells weremeasured with a confocal laser scanning microscope in tissuethat had been macerated by digestion with pectinase and in thicktangential sections. The lengths of early wood and late woodvessel elements were measured in tissues that had been maceratedby Franklin's method. The vessel elements and fusiform cambialcells varied considerably in length within individual samples.The mean length of early wood vessel elements corresponded tothat of fusiform cells in the dormant cambium but not in thereactivated cambium. Significant differences were observed betweenthe mean lengths of dormant and reactivated fusiform cambialcells, between those of reactivated fusiform cambial cells andearly wood vessel elements, between those of reactivated fusiformcambial cells and late wood vessel elements, and between thoseof early wood and late wood vessel elements. The frequency distributionsof lengths of cambial cells were bimodal and differed from thoseof vessel elements, which more closely resembled a normal distribution.The proportion of shorter lengths was higher in the reactivatedcambium than in the dormant cambium, the early wood and thelate wood vessel elements. Our results do not suppot the hypothesisthat the lengths of early wood vessel elements in ring-poroushardwoods change during differentiation. The similar rangesof recorded lengths suggest that short and long vessel elementsmight be derived directly from short and long cambial cells,respectively. Copyright 1999 Annals of Botany Company Kalopanax pictus, cambium, vessel element, confocal laser scanning microscopy, maceration, cell length.     

Seasonal variation in the ultrastructure of the cambium in young stems of willow (Salix viminalis) in relation to phenology

The growth period of Salix viminalis L. (clone 683) plants near Stockholm, Sweden, (59.5°N, 18.3°E) started in April with flowering and ended in October with abscission of the shoot tips. Cell divisions in the vascular cambium started almost two months before sprouting and ceased at about the same time as the elongation growth of the shoots. Phloem cells were apparently produced before flowering, while new xylem production started at the time of flushing. Cytodifferentiation in immature xylem continued until November. Thick-walled cells with protoplasm were observed adjacent to xylem mother cells in the cambium during the winter. The number of radially arranged cells in the cambial zone increased from 3–4 during dormancy to about 18 during the mitotic maximum in July. Seasonal variation was apparent in vacuolization, wall thickness and presence of storage material in the cells. Lipid bodies and protein bodies occurred in both fusiform and ray initials, while starch was observed in ray initials, ray cells and in the phloem. In September the ultrastructure of the cambium showed anatomical features characteristic for both active and dormant cells. Dictyosomes with vesicles and rough ER were present in thick-walled cells that contained lipid bodies and starch granules. Nuclear divisions in the cambium ended in October.     

毛白杨维管形成层带细胞超微结构的季节性变化

木材(次生木质部)是树木形成层细胞分化的产物,形成层的活动方式不仅影响木材的产量,而且影响木材的结构和性质.利用透射电子显微镜观察了生长在北京地区的毛白杨(Populus tomentosa Carr.)枝条形成层带细胞一个完整活动周期的超微结构变化.在木质部母细胞完全恢复活动之前,形成层纺锤状原始细胞的分裂和韧皮部细胞的分化已经开始.枝条上芽的展开和幼叶的生长可能决定了形成层带细胞的这种活动方式.透射电镜观察更清楚地揭示了树木形成层细胞在活动初期的分化特点.活动期形成层细胞中的大液泡在进入休眠期后逐渐分成许多小液泡分散在细胞质中.随着液泡融合逐渐消失的深色蛋白类物质又重新充满了大部分液泡.油滴和淀粉颗粒的年变化情况同液泡中的蛋白类物质基本相似.无论在活动期还是休眠期,形成层纺锤形细胞的质膜上都发现有许多可能与物质运输有关的小泡状内折.由核膜、内质网和高尔基体及其分泌小泡组成的细胞内膜系统,在形成层活动周期的不同阶段,其形态和分布明显不同,尤其在形成层细胞的恢复活动及其衍生木质部细胞次生壁的沉积过程中发挥着重要作用.整个活动周期中,形成层纺锤形细胞的径向壁都比弦向壁厚,处在休眠期的形成层带细胞,其径向壁与弦向壁的差别则更明显.形成层恢复活动时,径向壁上特别是与弦向壁相连的角隅处出现部分自溶现象.细胞壁特别是径向壁的变薄是形成层细胞恢复活动的重要特征.