AN INVESTIGATION OF THE PHYLOGENETIC AND ONTOGENETIC VARIABILITY OF SHOOT APICAL MERISTEMS IN THE CACTACEAE

The sizes, shapes and zonations of the shoot apical meristems of 22 species of cacti were examined. This family was chosen because of its great diversity of habits; the more primitive members are nonsucculent. leafy trees and more advanced members are highly specialized “leaf-less” stem-succulents. By combining these measurements with those already in the literature, a sample of almost 70 species was obtained. Apical meristems range in size from only 80 μm in diam in some species to as much as 1.500 μm in diam in others. The shape ranges from being flat to almost hemispherical. Despite the great range in size and shape of the apical meristems, or the range in the morphologies of the leaves and stems which are produced by the meristems. all apices had the usual zonation: tunica, central mother cells, peripheral zone, and pith-rib meristem. The sizes of each of the zones. expressed either as the number of cells per zone or expressed as a percentage of the whole apex. were highly variable. The variation in apical dimensions and zone sizes occurred both phylogenetically and ontogenetically. and this has been interpreted to indicate that the morphogenetic mechanisms which control apical size and zonation are easily modified, both during the development of individual plants and during the evolution of new species.     

香榧营养苗端细胞组织分区的超微结构观察

本文研究了榧树(Torreya grandis)成熟植株在季节生长中营养苗端的超微结构变化。各区域细胞的主要区别特征为:顶端原始细胞与亚顶端细胞相接的细胞壁较厚,液泡多分布于细胞游离面,质体中淀粉粒较小;亚顶端细胞壁较厚,液泡较大,质体中淀粉粒较大而多;周缘区细胞质体多不具淀粉粒,液泡也较小,胞间连丝丰富;肋状区细胞被大量的含淀粉质体及液泡占据了大部分空间,胞间连丝丰富。在季节变化的四个时期中,各区域细胞的亚显微结构特征亦不相同。休眠期各区细胞淀粉质体较发达,细胞壁较厚,液泡大;叶扩展期淀粉质体减少或消失;芽鳞形成期出现大量小液泡;新的顶芽形成期液泡增加,核糖体含量较高。讨论了各区域细胞核形态与其细胞活跃性的关系。     

AN ELECTRON MICROSCOPIC EVALUATION OF CYTOHISTOLOGICAL ZONATION IN THE SHOOT APICAL MERISTEM OF PINUS BANKSIANA

Shoot apical meristems of jack pine (Pinus banksiana) were examined by light and electron microscopy. Cytohistological zonation was evident when meristems were fixed in Craf IV, embedded in paraffin, and stained with Chlorazol Black E. When meristems were fixed for electron microscopy the cytoplasm of the apical initials and central mother cells each contained numerous lipid bodies and their nuclei contained little, if any, heterochromatin. The cytoplasm of the peripheral zone was rich in ribosomes. The nuclei of the peripheral zone and rib meristem were heterochromatic. Thus, the lack of heterochromatin in the nuclei and the dissolution of lipids in the cytoplasm of the apical initials and central mother cells appeared to contribute most to the organization and appearance (cytohistological zonation) of the shoot apex when standard histological techniques are used.     

IN VITRO DEVELOPMENT OF THE ISOLATED SHOOT APICAL MERISTEM OF ANGIOSPERMS

Development of complete plants was achieved from isolated shoot apical meristems of Nicotiana tabacum L., Daucus carota L., Nicotiana glauca Grah., Tropaeolum majus L., and Coleus blumei Benth. The explants consisted of only meristematic dome tissue with no visible leaf primordia. A simple nutrient medium composed of the Murashige and Skoog salt mixture, 100 mg/liter myo-inositol, 0.4 mg/liter thiamin-HCl, 1-2 mg/liter IAA, 30 g/liter sucrose, and 1% agar was adequate. Histologically there occurred principally tissue enlargement during the first 3-6 days, followed by appearance of bipolar organization in 6-9 days and formation of a well-defined root apex and initiation of first leaf primordium by 12 days.     

SIMULATIONS OF CELL DIMENSIONS IN SHOOT APICAL MERISTEMS: IMPLICATIONS CONCERNING ZONATE APICES

Surface areas, differential curvatures, volumes, and dimensions of cell-like units were described for various geometric shapes approaching the morphology of shoot apical meristems (SAM) in vascular plants. Geometric relationships are given in both graphic and dynamic simulations of shape. If the surface of the SAM is described by the revolution of a parabolic, hyperbolic, or circular function, then the greatest change in the surface areas or volumes of equally spaced transverse sections will occur in those regions showing the greatest differential in surface curvature. Subdivision of the SAM volume into cell-like units leads to quantitative expressions of changes in “cell” length to width and width to depth (aspect) ratios. Dependent upon the geometry of the SAM, differential expressions of aspect ratios may lead to a zonate pattern within the SAM corresponding to a central-mother-cell zone (CMC), a peripheral zone (PZ), and a pith-rib meristem (PRM). The boundary between the PRM and PZ, as seen in median longitudinal section, is a geometric consequence of the deployment of cells with aspect ratios best suited to occupy the entire SAM volume. The number of cell-like lineages, L, in the SAM may be expressed by the aspect ratio of cells in longitudinal section, n_x-y, such that for the PRM and PZ, L = k₁n_x-y^-5.38 and L = k₂n_x-y^3.18, respectively. Cellular patterns seen in the “corpus” of the SAM may not, therefore, be the a priori result of physiologically distinct populations of cells. Computer simulations are discussed within the data set derived from a study of the SAM of cacti.     

ZONATION IN THE APICAL CELL OF ZONARIA

Light and electron microscopic techniques were used to study the row of apical cells that form the meristem of the dictyotalean brown alga, Zonaria farlowii. A distinctly polar pattern occurs in the cells. Four ultrastructurally different cytoplasmic zones have been seen: an apical zone, a nuclear zone, a compound vacuolar zone, and a vegetative zone. The apical cell of Zonaria is a differentiating cytoplasmic unit where striking intracellular gradients occur.     

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.     

CYTOKININ-ELICITED FORMATION OF THE PITH-RIB MERISTEM AND OTHER EFFECTS OF GROWTH REGULATORS ON THE MORPHOGENESIS OF ECHINOCEREUS (CACTACEAE) SEEDLING SHOOT APICAL MERISTEMS

Shoot apical meristems of Echinocereus engelmannii have only a tunica-corpus organization at germination, but the corpus rapidly develops central mother cells, a peripheral zone and a pith-rib meristem. The manner in which nutrition, darkness and various growth regulators at several concentrations and in several combinations affect the development of zonation was examined by growing derooted seedlings on agar which contained the nutrients or growth regulators. Benzylaminopurine was able to elicit the formation of the pith-rib meristem in an otherwise non-zonate corpus. Also, the rate of leaf initiation was greatly increased. Gibberellic acid severely inhibited the formation of corpus zones but had little effect on leaf initiation. Indoleacetic acid had no effects other than mild inhibition of zonation and a slight retardation of leaf initiation. Abscisic acid was strongly inhibitory. Sucrose only slightly increased the rate of leaf formation and did not affect apex size or zonation. To more closely examine the cytokinin-induced effects on the apical meristem, several growth regulators were applied in combination with the most effective concentration of cytokinin. Certain combinations were able to interfere with several of the cytokinin-induced responses, while other cytokinin-induced responses occurred even in the presence of high concentrations of these other growth regulators. Leaf initiation and meristem morphogenesis appeared to be remarkably stable and insensitive to the presence of most hormones except cytokinin and gibberellin.     

DEVELOPMENT OF VACUOLES AND LIPID BODIES IN APICAL MERISTEMS OF PINUS BANKSIANA

Shoot apical meristems of jack pine were examined weekly during the first 8 weeks post-germination with light and electron microscopy. Most of the storage lipids were utilized by the end of the 2nd week. A few lipid bodies, possibly high in phospholipid content, remained in the apical initials and central mother cells and, during the 3rd week, gave rise to vacuoles via lamellar or myelin-like structures which were first seen on their periphery. The inter-lamellar spaces enlarged and eventually a vacuole was formed. At 5 weeks, elongate and spherical osmiophilic inclusions, presumably lipid, were found in the endoplasmic reticulum. Lipid bodies, visible with light microscopy, began to accumulate in the apical initials and central mother cells in the 6th week.     

SHOOT APICAL MERISTEMS AND MUTATION: FIXATION OF SELECTIVELY NEUTRAL CELL GENOTYPES

Shoot apical meristems are interpreted as either structured, that is having a permanent set of apical initials, or stochastic, having apical initials which represent “... momentary representatives of the continuous meristematic residue at the apex of the relevant layer or zone” (Newman, 1965). The two main parameters of stochastic growth are the average number of apical initials (α) and the number of mitotic cycles (r) of the initials and their daughter cells prior to the random selection of subsequent initials. Mathematical analysis and computer simulation studies of stochastic growth have shown that if one starts with 1 mutant initial and α-1 nonmutant initials, eventually a mosaic plant results. The frequency of shoot apices composed of mutant cells is 1/α and the frequency of shoot apices composed of only nonmutant cells is (1 – α)/α. These asymptotics are only attained after considerable growth, thus mericlinal chimeras can persist for many nodes and give the appearance that a permanent set of initials is present.     

芸薹属植物发育过程中顶端结构的解剖学研究

本文采用解剖学方法研究花椰菜、青花菜、结球甘蓝和大白菜在生长发育过程中顶端分生组织结构的变化及之间存在的差异。结果显示它们的顶端分生组织结构都是由最初幼苗的原套-原体结构逐渐发育到过渡型分区结构、典型化五个分区结构,至开始进入生殖生长时期的四个分区结构(形成层状细胞区消失)。四种植物在进入生殖生长后,顶端分生组织细胞行为不同:大白菜和甘蓝顶端亚外套两侧细胞分裂分化形成顶生叶原基,在顶生叶原基内侧的细胞将进行分裂产生花序侧枝原基。花椰菜和青花菜顶端亚外套两侧细胞分裂形成花序分生组织,花序分生组织增生即为花球体;内部解剖结构表现为分生组织不断分裂增多的过程。这些结果为研究花序表型发生的解剖学本质及分子生物学研究分生组织发育方向奠定了基础。     

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.     

CYTOKININ- AND GIBBERELLIC ACID-INDUCED EFFECTS ON THE STRUCTURE AND METABOLISM OF SHOOT APICAL MERISTEMS IN OPUNTIA POLYACANTHA (CACTACEAE)

The dormant axillary buds of Opuntia polyacantha can be activated by either cytokinins or gibberellic acid. Under the influence of benzylaminopurine (BAP), the axillary bud meristem increases greatly in size and becomes mitotically active. The primordia produced by the meristem develop as normal photosynthetic leaves. Gibberellic acid (GA) also causes the meristem to become mitotically active, but the meristem does not increase in size. The primordia produced under the influence of GA develop as normal cactus spines. Leaf-producing meristems and spine-producing meristems have the same zonation, despite the differences in size. The meristems are composed of a uniseriate tunica, a central mother cell zone, peripheral zone, and a pith rib meristem. The mitotic activity of each of the zones in the leaf-producing meristem differs significantly from the mitotic activity of the corresponding zones in the spine-producing meristem.     

THE SHOOT APICAL ONTOGENY OF THE PICEA ABIES SEEDLING. III. SOME AGE-RELATED ASPECTS OF MORPHOGENESIS

We have extended our previous analyses of growth in shoot apices of Picea abies seedlings. Quantification of apical dome volume changes, more detailed analysis of the subapical caulis profile, and of the vertical distance from the base of the dome to the nth primordium, all as functions of age, revealed the dynamics of various growth variables. As seedlings age from 10 to 136 days, apical dome volume increases about 30-fold, plastochron duration declines from 31 h to about 5 h, height of primordial internodes declines from near 10 μm, to only 3.5 μm, and the caulis assumes a distinct neck-and-shoulder profile. Relative volume growth rates for the apical dome as a whole are about twice the base-of-dome values and decline from 36% to 21% per day as age increases to 136 days. Relative growth rates (radial, vertical, and volume) in the caulis change in a complex manner with both plant age and internode number. We also computed the total volume of tissue generated by an apical dome per day including that part invested in increased dome volume. The investment (ϕ) ratio is greater than 20% in the 10-day dome, but declines rapidly to become negative after 136 days. The ϕ ratio controls apical dome volume and hence augurs future growth yield.     

除草剂拿草特对菟丝子幼苗茎尖分生区有丝分裂的影响

酰胺类除草剂拿草特对菟丝子幼苗生长具有强烈的抑制作用。采用石蜡切片法和细胞扫描分光光度法,通过测定对照及处理幼苗茎尖分生区的细胞分裂指数和核DNA含量,发现拿草特对菟丝子幼苗生长的抑制是由于它对分生区细胞有丝分裂的双重抑制作用:间期抑制和中期抑制。即它既可阻止有丝分裂的启动,又可象秋水仙素一样通过影响纺锤丝的功能而将细胞分裂阻止于中期,引起分生区细胞的多倍体化和径向膨大。这种生长受到抑制且茎尖粗肿的幼苗不能够在农作物上寄生。     

THE SHOOT APICAL ONTOGENY OF THE PICEA ABIES SEEDLING. IV. PROTOXYLEM INITIATION AND AGE OF INTERNODES

In Picea abies seedlings the distance below the base of the shoot apical dome to the first protoxylem (px) to be differentiated in the internodes beneath is a linear function of apical dome basal diameter. By using mathematical relations derived in earlier papers of this series, we computed the morphogenic age in plastochrons and the chronometric age in days of the internode in which px is first differentiated (n_px). As the seedlings age from 30 to 140 days, the distance from the base of the apical dome to px increases from 186 to 295 μm, the n of n_px increases from 16 to 53, but the chronometric age of n_px remains within the range of 10 to 12 days. Protoxylem differentiation in young internodes is, therefore, more closely related to chronometrie age than it is to morphogenic age or to distance from the base of the shoot apical dome.