CELL DIVISION STUDIES OF THE SHOOT APEX OF DATURA STRAMONIUM DURING TRANSITION TO FLOWERING

Seedlings of Datura stramonium L., although not photoperiodically sensitive, are useful for floral transition studies when raised in a growth chamber at a constant temperature of 25 C with a photoperiod of 8 hr of light (1,600-2,000 ft-c) and 16 hr of darkness. A terminal flower is formed after the seventh or eighth leaf primordium is produced. A constant rate of leaf initiation up to the time of flowering enables specific apical stages to be obtained and studied. Changes in the mitotic index, substantiated with calculated rates of cell division (measured by the accumulation of metaphases following treatment with colchicine) were studied in shoot apical zones during transition to flowering. Fluctuations in the mitotic index of each zone in the vegetative and transition apex with respect to apical stage as well as time of day were not statistically significant. The mitotic index of the summit zone of the vegetative apex was significantly lower than in the other zones whose mitotic indices were not significantly different from one another. During floral transition the mitotic index of the summit zone as well as the central zone (just below the summit zone) significantly increased while no significant changes were detected in the flank zones. It was shown that the mitotic index could be considered representative of the rates of cell division in Datura.     

THE EARLY ACTION OF THE FLORAL STIMULUS ON MITOTIC ACTIVITY AND DNA SYNTHESIS IN THE APICAL MERISTEM OF XANTHIUM STRUMARIUM

Vegetative plants of Xanthium strumarium (a short-day species) were induced to flower by exposure to a single 16-hr long night. By cutting off the induced leaf (half-expanded leaf) at various times, it was established that, by 8 hr after the end of the long night, a sufficient amount of floral stimulus had reached the meristem to induce a flowering response. The following sequence of events occurred in both the peripheral and central zones of the apical meristem of induced plants: 1) a rise in the mitotic index beginning at 28 hr after the end of the long night and culminating at 36 and 56 hr; 2) a stimulation of DNA synthesis starting at 32–36 hr and reaching a maximum at 60 hr; 3) an increase in nucleolus diameter starting at 32 hr. The cell population in the meristems of both vegetative and induced plants displayed a similar distribution, with about 80 % of the nuclei with the 2C amount of DNA. The comparison of the kinetic data concerning the mitotic index and DNA synthesis indicated that one of the early effects of the floral stimulus in the peripheral and central zones was the release in mitosis of cells whose nuclei were in the postsynthetic (G₂) phase of the mitotic cycle. In the pith-rib meristem, the following events were recorded: 1) a stimulation of DNA synthesis starting at 20 hr; 2) a rise of the mitotic index beginning at 28 hr; 3) the vacuolation and elongation of cells starting at 48 hr. All these events occurred well before the initiation of bract and flower primordia, which began at 96 and 136 hr, respectively. Neither stimulation of mitotic activity nor flowering occurred in the meristems of plants subjected to a long night interrupted at its midpoint by a 5-min light break. The results are discussed in relation to the early events which are known to occur in the meristems of other photoperiodic species in transition to flowering.     

INFLORESCENCE DEVELOPMENT IN BRASSICA CAMPESTRIS L.

Vegetative seedlings of the Ceres strain Brassica campestris L., a quantitative, long-day plant, were induced to flower by exposure to a 16-hr, long-day cycle. Cytohistological and cytohistochemical changes associated with inflorescence development were examined. Developing shoot apices were classified in vegetative, transitional, and reproductive stages. The vegetative apex possessed a biseriate tunica, central zone, peripheral zone and pith-rib meristem. The transitional stage at 48 hr was marked by an increase in size and by a stratification of the upper cell layers of the shoot apex with a concurrent decrease of apical cytohistochemical zonation. The reproductive stage was initiated at 58 hr by periclinal cell divisions in the 3rd and 4th cell layers of the flank region. Cytohistochemical zonation in the vegetative apical meristem was restored in the floral apex. An “intermediate developmental” phase was not observed between the vegetative and reproductive stage.     

VARIATION IN CELL-CYCLE TIME AND NUCLEAR DNA CONTENT IN THE APICAL MERISTEM OF HELIANTHUS ANNUUS L. DURING THE TRANSITION TO FLOWERING

The mitotic cycle in the apical meristem of Helianthus annuus L. has been investigated during the transition to flowering. Towards the end of the strictly vegetative phase 8 days after sowing the average cell-cycle time, measured by colchicine-induced metaphase accumulation, was 37 hr in the peripheral zone, 83 hr in the central zone and 118 hr in the rib meristem. By Day 12 the cycle had shortened in all zones. By the time of floral initiation on Day 16 the cycle time had returned to its original value in the peripheral zone and the rib meristem, while in the central zone it continued to shorten to 33 hr, approaching the cycle time of the peripheral zone. Cytophotometric measurements of nuclear DNA showed that mitotic activation of the central zone was not associated with any reduction in the proportion of nuclei with a 4 C DNA content. It was calculated that the spatial and temporal variation in cell-cycle time was mainly a function of the length of the G₁/G₀ phase which lasted about 19 hr in the peripheral zone, 82 hr in the rib meristem, and declined from 55 to 21 hr in the central zone.     

The pattern of histone H4 expression in the tomato shoot apex changes during development

Two histone H4 cDNA clones were isolated from a tomato (Lycopersicon esculentum Mill.) shoot-tip cDNA library using a heterologous probe from barley (Hordeum vulgare L.). Both cDNAs, which are 81% identical in the coding region, are polyadenylated and belong to a small gene family in the tomato genome. Histone H4 message is abundant in young tissues and rare in older tissues. In the shoot apical meristem, the distribution of H4-expressing cells changes during development. In a juvenile vegetative apex, H4 message is detectable in the central region and the peripheral parts of the meristem. In a mature vegetative apical meristem, H4-expressing cells are localized in the peripheral zone extending into the provascular strands and the rib meristem whereas the central zone is almost devoid of H4 mRNA. After floral transition, H4 mRNA is found throughout the floral meristem, indicating a second change in the pattern of H4 expression. The observed changes in H4 expression are indicative of changes in the distribution of mitotic activity in the shoot apical meristem during plant development. In addition, H4-expressing cells were found to occur frequently in clusters, which may indicate a partial synchronization of cell divisions in the shoot apex.     

HISTOCHEMICAL STUDY OF ENZYME ACTIVITY IN THE SHOOT APICAL MERISTEM OF BRASSICA CAMPESTRIS DURING TRANSITION TO FLOWERING. IV. GLUCOSE-6-PHOSPHATASE

Glucose-6-phosphatase (G6P) activity was determined in fresh-frozen, cryostat sections in the shoot apical meristem of Brassica campestris L. Enzymatic activity was differentially distributed in a zonate pattern in the vegetative meristem, but not in the transition and floral meristem. Vegetative apices showed a heterogenous localization with the highest activity in the central zone and the pith-rib meristem zone. At the early transition stage of development, G6P activity in the peripheral zone increased slightly. At the late transitional (prefloral) stage, G6P activity was not localized within the peripheral zone in island-like areas of activity. This is the first demonstration of G6P in shoot apical meristem at the vegetative, transition, and floral stage. The results indicate that G6P activity 1) is an accompanying event of evocation, but 2) does not mark incipient floral primordia. G6P may play an important role in the maintenance of glucose-6-phosphate homeostasis in an evoked shoot apical meristem.     

Structure and Cytogenesis of the Shoot Apex of Syringa oblata var. affinis Lingelsh

The structure, growth and mitotic activity of 211 shoot apices of developing sprouts of Syringa oblata var. affinis Lingelsh. in longitudinal sections and 67 in transverse sections have been studied with the view to understanding the nature of zonation patterns and cytogenesis of the apical meristems during a double plastochron. The external morphology and the anatomical structure of the apices in 4 plastochronic stage-early, middle, late Ⅰ and late Ⅱ stages are described. In the shoot apices examined, especially those at late plastochronic stage, the following zones may be delimited: Zone of tunica initials, zone of corpus initials, peripheral zone and zone of rib meristem. The location and orientation of mitotic figures observed in longisections of the apices in 4 plastochronic stages are plotted in diagrams and the mitotic frequency has been calculated. Information obtained from these investigations reveals that the tunica and corpus inititals constitute an active region of the apex, but their mitotic activity changes periodically within the double plastochron. In the middle plastochronic stage when the apex is at its minimal area and the cells of peripheral zone and rib meristem zone have been completely transformed into constituent parts of foliar primordia and the subjacent tissues of the stem and the pith mother cells respectively, the mitotic frequency of the initials is at its maximium and its intensity of mitotic activity is not much lower than that of any other meristematic zone at any stage. When the apical dome is reformed by the activity of these initials in late plastochronic stages, the mitotic frequency of the initials gradually drops and the region of high mitotic frequency shifts to the flank of the apex, the peripheral zone. Anticlinal divisions are predominant in this zone. On the other hand, those cells directly left behind by the corpus initials, which constitute the rib meristem, are vacuolated and marked by the pre- dominance of transverse divisions. Thus the entire zonation pattern reappears. In the next early plastochronic stage, the mitotic frequency of the tunica and corpus initials drops to its mimimium, but other regions of the apex still maintain a high mitotic frequency. It may be concluded that the tunica and corpus initials form a cytogenerative center of the shoot, and the cytohistological zonation is actually a result of the fact that different regions of apical meristems are different in mitotic activety, different in state of cell differentiation and different in their function in morphogenesis.     

CHANGES IN GLYCERALDEHYDE 3-PHOSPHATE DEHYDROGENASE ACTIVITY IN SHOOT APICAL MERISTEMS OF BRASSICA CAMPESTRIS DURING TRANSITION TO FLOWERING

An assay system has been developed for the histochemical determination of glyceraldehyde 3-phosphate dehydrogenase (G3PD) activity to indicate glycolytic pathway capacity during evocation in the shoot apical meristem of Brassica campestris L. G3PD activity was differentially distributed in a zonate pattern within the meristems at the vegetative, the transition, and the floral stages. The activity of G3PD changed in all apical zones of evoked apices, but especially in the central and the peripheral zones of apices at the prefloral stage. In the prefloral stage of development heavy enzyme activity was localized in island-like areas within the peripheral zone. These results indicate that 1) the capacity of glycolysis fluctuates during evocation, 2) during floral evocation the capacity of the glycolytic pathway parallels the capacity of the citric acid cycle and the electron transport system only at the prefloral stage, and 3) G3PD activity marks incipient floral primordia. It is proposed that the enzymic marking of an incipient floral primordium indicates the end of evocation in Brassica.     

CONSTANCY OF RELATIVE VOLUMES OF ZONES IN SHOOT APICAL MERISTEMS IN CACTACEAE: IMPLICATIONS CONCERNING MERISTEM SIZE,SHAPE, AND METABOLISM

The volumes and dimensions of shoot apical meristem zones were determined for 21 species in 10 genera of the Cactaceae. If the volumes of the central-mother-cell zone (CMCZ), the peripheral zone (PZ), and the pith-rib meristem (PRM) are expressed as percentages of the total volume of the corpus, then they are remarkably constant (CMCZ = 11.1% of the volume of the corpus, PZ = 61.2% and PRM = 27.1%) regardless of the relative size of the apical meristems. The relative volume of the tunica decreases, however, as the whole apex increases. The zones have variable shapes, and whereas the PZ and PRM are always trapezoidal in longitudinal section, in apices with large volumes, these zones have a flatter, more flaring shape than they have in smaller apices. Thus large apices are flatter and less hemispherical than are small apices. The CMCZ, however, maintains a constant shape and is always relatively hemispherical regardless of its volume. A hypothesis that relates all of these shape and volume relationships as an integrated growth sequence is proposed.     

香榧营养苗端的结构及淀粉动态的研究

香榧（Torreya grandis Fort ex Lindl)成熟植株营养芽的季节生长可分四个时期,休眠期,叶扩展期,芽鳞形成期和新的顶芽形成期,在整个生长周期中,苗端始终保持一定的分区形式,但各区繁简在不同发育阶段不尽相同,淀粉在苗端的分布及消长也具有分区特性,与细胞组织学分区完全一致。     

CHANGE IN EPILOBIUM PHYLLOTAXY DURING REPRODUCTIVE TRANSITION

The ontogeny of Epilobium hirsutum grown under natural summer photoperiod in a glasshouse was divided into vegetative, early transitional, transitional, and floral stages. Bijugate phyllotaxy, common to both the vegetative and early transitional stages, is transformed into spiral phyllotaxy during the transitional stage by an initial change in the divergence angle of a single primordium inserted at a unique level on the shoot. Leaf primordia subsequently are inserted in a spiral arrangement in the indeterminate floral shoot apex. The early transitional shoot apical meristem is about 1.5 times the volume of the vegetative meristem but expands at about two-thirds the relative plastochron rate of volume increment of the vegetative meristem. There are progressive decreases in the plastochron and relative plastochron rates of radial and vertical shoot growth through ontogeny. Relative chronological rates of shoot growth, however, are not altered during ontogeny. Spiral transformation results from changes in the relative points of insertion of leaf primordia on the shoot meristem. These changes are accompanied by an increased rate of primordia initiation on a more circular shoot meristem. The change in phyllotaxy during ontogeny is similar to that which was artificially induced by chemical modification of auxin concentration gradients in the shoot apex, with the additional feature that there is an initial increase in the volume of the shoot meristem prior to the natural spiral transformation. Size of the shoot apical meristem, however, appears to have little influence on Epilobium phyllotaxy; but the geometric shape of the meristem is well correlated with bijugate to spiral transformations. This suggests that geometric parameters of the shoot meristem should be considered in theoretical models of phyllotaxy.     

TENDRILS OF PASSIFLORA FOETIDA: HISTOGENESIS AND MORPHOLOGY

Passiflora foetida bears an unbranched tendril, one or two laterally situated flowers, and one accessory vegetative bud in the axil of each leaf. The vegetative shoot apex has a single-layered tunica and an inner corpus. The degree of stratification in the peripheral meristem, the discreteness of the central meristem, and its centric and acentric position in the shoot apex are important plastochronic features. The procambium of the lateral leaf trace is close to the site of stipule initiation. The main axillary bud differentiates at the second node below the shoot apex. Adaxial to the bud 1–3 layers of cells form a shell-zone delimiting the bud meristem from the surrounding cells. A group of cells of the bud meristem adjacent to the axis later differentiates as an accessory bud. A second accessory bud also develops from the main bud opposite the previous one. A bud complex then consists of two laterally placed accessory bud primordia and a centrally-situated tendril bud primordium. The two accessory bud primordia differentiate into floral branches. During this development the initiation of a third vegetative accessory bud occurs on the axis just above the insertion of the tendril. This accessory bud develops into a vegetative branch and does not arise from the tissue of the tendril and adjacent two floral buds. The trace of the tendril bud consists of two procambial strands. There is a single strand for the floral branch trace. The tendril primordium grows by marked meristematic activity of its apical region and general intercalary growth.     

The shoot apical meristem of Sinapis alba L. expands its central symplasmic field during the floral transition

The shoot apical meristem (SAM) is functionally subdivided into zones with distinct tasks. During vegetative growth the peripheral zone of the meristem gives rise to leaf primordia that develop into dorsiventral leaves under the influence of signals from the central zone. During the floral transition the function of the SAM is altered and its peripheral zone starts to form floral structures in a specific pattern. This requires alterations in the signal networks that coordinate the activities of the peripheral and central zone of the SAM. These signal networks are partly housed in the symplasmic space of the SAM. Dye-coupling experiments demonstrate that in the superficial layer of the Sinapis alba meristem this space is radially subdivided. The cells of the central zone are coupled into a symplasmic field, which is shielded from the peripheral zone by the positional closing of plasmodesmata. In the vegetative meristems, most of these central symplasmic fields have a triangular geometry and are relatively small in size. Plants that are induced to flower by exposure to a single long day alter the geometry as well as the size of their central symplasmic field. After two subsequent days under short photoperiod the central symplasmic fields exhibit a circular form. Simultaneously, their size strongly increases both in an absolute sense and relative to the enlarging meristem. The geometric change in the fields is hypothesized to be due to recruitment of extra initial cells, required to support the increase in phyllotactic complexity. The proportional increase in field size is interpreted as an adjustment in the balance between the central and peripheral zone of the SAM, accompanying the shift from leaf production to flower formation.     

INITIATION AND EARLY DEVELOPMENT OF THE INFLORESCENCE IN PINEAPPLE (ANANAS COMOSUS, ‘SMOOTH CAYENNE') TREATED WITH ACETYLENE

Pineapple plants ‘Smooth Cayenne’ were made to flower by treatment with acetylene. The organization of the vegetative shoot apex is similar to that of many investigated angiosperms in that it shows a zonate pattern, viz., apical zone, peripheral zone, and central-core rib meristem. The latter zone is weakly developed. Cytological changes at the shoot apex occur as early as 3 days after treatment; these involve nuclear changes and an increase in ribonucleic acid (RNA) in the cytoplasm of cells of the apical zone. A marked increase in the height of the apex occurs by the 9th day; this is preceded by rib meristem activity in the central core. All component parts of the inflorescence are present and in various stages of development by the 21st day at which time vegetative scales and “crown” leaves are initiated.     

A HISTOCHEMICAL STUDY OF THE SEEDLING SHOOT APICAL MERISTEM OF PINUS LAMBERTIANA

Vernalized seeds of Pinus lambertiana were scarified and planted in perlite. At 5, 8, 10, 13 and 16 days after planting, seedlings were selected for morphological examination and histochemical study. The shoot apical meristem consisted of a relatively homogeneous population of cells at 5 days. Cytohistological zonation was observed in the meristem by the eighth day and needle primordia initiation began at this time. Acid phosphatase (AP) activity was high in the extreme tip of the apex at 5 days. At 8 days AP activity was intense in the peripheral zone but weak in the apical initial and central mother cell zones. The apical meristem of the 10–16-day-old seedlings exhibited high AP activity in the peripheral zone only during the early stages of needle primordia initiation. The distribution of cytoplasmic and nuclear protein-bound SH was correlated with cytohistological zonation. Protein-bound SH was distributed relatively uniformly at 5 days, but by the eighth day the 4 cytohistological zones contained differential quantities. Succinic dehydrogenase (SD) activity was observed throughout the apex at 5 days, but by the eighth day the apical initial and central mother cell zones exhibited differentially greater levels of SD activity. Irradiation with 500 R of X-rays at 7 days after planting completely inhibited needle primordia initiation and disrupted the cytohistological zonation of the apex. Correlated with the inhibition of needle primordia initiation was the loss of SD activity in the apical initial and central mother cell zones. Irradiation also resulted in the gradual loss of protein-bound SH from the cytoplasm of the apical initial, central mother cell and peripheral zone.     

Les critères nucléaires d'une dormance vraie et totale dans le point végétatif du Fraxinus excelsior L.

Summary The cytohistological criteria for the vegetative shoot apex dormancy in Fraxinus excelsior L. have been quantitatively established with the aid of 3 techniques: historadiography after incorporation of [³H]thymidine, mitotic index and nuclear cytophotometry by the two wavelength method. Nuclear DNA content, mitotic activity and DNA synthesis were compared in 3 different zones(apical zone, lateralzone, rib meristem) of the dormant and non-dormant apices. The periodical break in morphogenetic activity, in contrast to the vegetative period (April to July), is characterized by the absence of zonation and by the fact that meristematic cells remain in the G1 phase of the mitotic cycle. In Fraxinus excelsior L., the meristem dormancy is complete (no DNA synthesis, no mitotic activity and no DNA content greater than 2C).

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Abréviations employées ZA zone apicale axiale du point végétatif - ZL zone latérale ou anneau initial - mm meristème médullaire - UA unité arbitraire de quantité de DNA     

QUANTITATIVE STUDIES OF THE VEGETATIVE SHOOT APEX OF EQUISETUM SCIRPOIDES

Equisetum scirpoides Michx., propagated from a single clone, was grown in a controlled growth chamber at 24 ± 1 C under a photoperiod of 16 hr light/8 hr darkness. The apical cell of aerial vegetative shoots gives rise to derivatives (merophytes) in a helical sequence. Each newly formed merophyte divides anticlinally to form two superposed cells that are parallel to a lateral face of the apical cell. Radial longitudinal divisions then take place in the two superposed cells. Shoot tips were fixed every 2 hr for 24 hr to determine the mitotic index of the apical cell, six subjacent cells, and the remaining cells above the level of leaf initiation. Average mitotic indices for the 24-hr period were 3.9%, 3.9%, and 7.0%, respectively. The results indicate that the apical cell is quite active mitotically; there was no clear evidence of endopolyploidy in cells of the shoot apex, young leaves or in the developing cortex, based upon cytophotometric measurements of DNA content.     

ONTOGENY OF THE INFLORESCENCE OF SAURURUS CERNUUS (SAURURACEAE)

The spicate inflorescence of Saururus cernuus L. (Saururaceae) results from the activity of an inflorescence apical meristem which produces 200–300 primordia in acropetal succession. The inflorescence apex arises by conversion of the terminal vegetative apex. During transition the apical meristem increases greatly in height and width and changes its cellular configuration from one of tunica-corpus to one of mantle (with two tunica layers) and core. Primordia are initiated by periclinal divisions in the subsurface layer. These are “common” primordia, each of which subsequently divides to produce a floral apex above and a bract primordium below. The bract later elongates so that the flower appears borne on the bract. All common primordia are formed by the time the inflorescence is about 4.4 mm long; the apical meristem ceases activity at this stage. As cessation approaches, cell divisions become rare in the apical meristem, and height and width of the meristem above the primordia diminish, as primordia continue to be initiated on the flanks. Cell differentiation proceeds acropetally into the apical meristem and reaches the summital tunica layers last of all. Solitary bracts are initiated just before apical cessation, but no imperfect or ebracteate flowers are produced in Saururus. The final event of meristem activity is hair formation by individual cells of the tunica at the summit, a feature not previously reported for apical meristems.     

AN INVESTIGATION OF THE MORPHOGENETIC MECHANISMS WHICH CONTROL THE DEVELOPMENT OF ZONATION IN SEEDLING SHOOT APICAL MERISTEMS

The development of zonation in the shoot apical meristems of 28 species of cacti was examined. At germination the embryonic apex may have one of three types of organization: 1) tunica/corpus; 2) tunica/central mother cells/corpus; 3) tunica/central mother cells/peripheral zone/pith-rib meristem. Apices of the third type have complete zonation and undergo little or no further structural development. Both of the other types develop the missing zones. First, the apices become mitotically active, and peripheral zone characters develop in the outer corpus. Simultaneously, or slightly later, the central mother cells differentiate if they are not yet present. The final step is the formation of the pith-rib meristem. The sequence of appearance of the zones was constant throughout all species examined, but the time of appearance of only one zone could be correlated with any other morphogenetic process: the development of peripheral zone characteristics in the outer corpus occurs with or before the beginning of leaf production. The development of zonation appears to be independent of apical size, shape, or age, either real age or plastochron age. This has been interpreted to indicate that the metabolic mechanism controlling the development of zonation in shoot apical meristems is largely autonomous and independent of other aspects of morphogenesis occurring in the seedling. Also, the fact that leaf initiation and shoot axis production can both occur before the development of either the central mother cells or the pith-rib meristem indicates that neither of these zones is essential for these two apical morphogenetic activities.     

Location of cell cycle changes in relation to morphological changes in the shoot apex ofSilene coeli-rosa immediately before sepal initiation

Summary Changes in morphology, the mitotic index and the proportions of cells in G₁ and G₂ were measured in shoot meristems ofSilene coeli-rosa immediately before floral morphogenesis in order to determine whether the known changes to the cell cycle at this time are restricted to a particular region of the apex. Twenty-eight day-old plants were given either 7 long days (LD) plus 2 short days (SD) (day 8 of the LD treatment) or 9 SD [day 8 of the SD control (SDC) treatment]. Plants were sampled on day 8 every 2 h for 12 h and the various cell cycle measurements were performed on sections of the apical meristem. In the inductive LD treatment there was a peak in the mitotic index at 13.00 h and, possibly, the start of another at 19.00 h. At 21.00 h all meristems in this treatment initiated sepals. The mitotic activity at 13.00 and 19.00 h in the LD treatment was a result of significant increases in the mitotic index in the axial, lateral and central sub-axial areas of the apex compared with the corresponding zones in the SDC treatment. At 13.00 h of day 8, 80% of cells were in G₂ phase in the axial region in the LD treatment whilst 85% of cells were in G₁ in the axial zone in the SDC treatment. In the other zones significantly more cells were in G₂ in the LD compared with the SDC treatment as was the case at 19.00 h although not to the same extent as the axial zone at 13.00 h. Thus these data emphasize, for the first time, the mitotic activation and predominance of the G₂ population of cells particularly in the axial zone of shoot meristems in the LD treatment. These data are discussed in relation to the synchronisation of cell division which could occur in the prefloral shoot meristem at this time, affecting each shoot apical zone.Abbreviations LD long day - SD short day - SDC short day control