Ficus rubiginosa 'variegata', a chlorophyll-deficient chimera with mosaic patterns created by cell divisions from the outer meristematic layer

BACKGROUND AND AIMS: Sections leaves of Ficus rubiginosa 'Variegata' show that it is a chimera with a chlorophyll deficiency in the second layer of the leaf meristem (GWG structure). Like other Ficus species, it has a multiseriate epidermis on the adaxial and abaxial sides of the leaf, formed by periclinal cell divisions as well as anticlinal divisions. The upper and lower laminae of the leaf often exhibit small dark and light green patches of tissue overlying internal leaf tissue. METHODS: The distribution of chlorophyll in transverse sections of typical leaves was determined by fluorescence microscopy. KEY RESULTS: Patches of dark and light green tissue which arise in the otherwise colourless palisade and spongy mesophyll tissue in the entire leaf are due to further cell divisions arising from the bundle sheath which is associated with major vascular bundles or from the green multiseriate epidermis. Leaves produced in winter exhibit more patches of green tissue than leaves which expand in mid-summer. Many leaves produced in summer have no spotting and appear like a typical GWG chimera. There is a strong relationship between the number of patches on the adaxial side of leaves and the number on the abaxial side, showing that the cell division in upper and lower layers of leaves is strongly coordinated. In both winter and summer, there are fewer patches on the abaxial side of leaves compared with the adaxial side, indicating that periclinal and anticlinal cell divisions from the outer meristematic layer are less frequent in the lower layers of leaf tissue. Most of the patches are small (<1 mm in longest dimension) and thus the cell divisions which form them occur late in leaf development. Leaves which exhibit large patches generally have them on both sides of the leaves. CONCLUSION: In this cultivar, the outer meristematic layer appears to form vascular bundle sheaths and associated internal leaf tissue in the entire leaf lamina.     

THE DEVELOPMENT OF APOSPOROUS GAMETOPHYTES AND REGENERATED SPOROPHYTES FROM EPIDERMAL CELLS OF EXCISED FERN LEAVES: AN ANATOMICAL STUDY

Three different types of outgrowths develop from epidermal cells of excised juvenile leaves of Microgramma vacciniifolia: aposporous gametophytes, intermediates, and regenerated sporophytic plantlets. The gametophytes and intermediates arise from derivatives of epidermal cell divisions which are developed to the exterior of the leaf surface, whereas the sporophytic regenerants originate from derivatives produced by cell divisions to the interior of the leaf. Anatomical observations of excised leaves grown in vitro demonstrate that only the epidermal cells are stimulated to divide and give rise to the various types of outgrowths. Incorporation of tritiated thymidine by the nuclei of leaf epidermal cells gives further evidence for the metabolic activity of these cells.     

Cellular basis of developmental plasticity observed in heterophyllous leaf formation of Ludwigia arcuata (Onagraceae)

When heterophyllous plants of Ludwigia arcuata Walt. (Onagraceae) were transferred from aerial condition to submergence, young developing leaves were matured into leaves with intermediate shape between aerial-type and submerged-type, showing spatulate shape (spoon-shaped). This change was also induced by the exposure of plants to ethylene. On the other hand, when the plants were transferred from submergence to aerial conditions, young developing leaves were matured into intermediate-type leaves with elliptic shape (spearhead shape). Anatomical analysis revealed that the formation of spatulate leaf was caused by the reduction of the number of epidermal cells aligned in the leaf transverse direction in the basal region of the leaf while the tip regions remained as before and did not respond to this treatment. During development, the ethylene-induced spatulate leaves showed that three types of alterations in epidermal cell division were involved in this process. Changes in the distribution of cell divisions in leaf lamina were detected by the first day of ethylene exposure, and changes in the orientation of cell division planes were detected by the second day. However, changes in the number of cells aligned in the leaf transverse direction were not detected by this time. Three days after ethylene exposure, frequency of cell divisions changed, and by the time changes of cell numbers aligned in the leaf transverse direction were observed. Thus, the formation of intermediate-type leaves in L. arcuata was ascribed to the alterations of cell division patterns which was induced by ethylene.     

不同生境下十七种藓类植物叶的比较解剖学

使用石蜡切片法,对不同生境下的17种藓类植物的叶片进行了解剖观察和比较分析,结果表明不同种类的藓类植物在中肋导水主细胞的有无、厚壁细胞是否分化、中肋细胞层数及细胞密度、叶片细胞层数、叶表附属物、叶片细胞密度等方面存在显著差异。藓类植物叶的解剖结构具有生态适应意义,旱生环境下的藓类植物,叶片细胞胞壁具不同程度的增厚,有些藓类植物叶片具附属结构,藓类植物中肋的有无,反映了对水分吸收和运输方式的不同。例如,荫湿生环境下的羽枝青藓Brachythecium plumosum,其中肋细胞胞壁较薄,无导水主细胞和副细胞的分化,也没有厚壁细胞分化,能够在阴湿环境下吸收水分和养分;钝叶匍灯藓Plagiom niumrostratum具有与旱生藓类植物相似的中肋结构,叶片较厚,中肋具导水主细胞,中肋背面具厚壁细胞,这些特点使该种藓类植物能够分布于间隙性干旱胁迫的环境中;水灰藓Hygro-hypnum luridum叶片纤细柔弱,仅1层细胞,细胞胞壁薄,叶表无附属结构,中肋细胞层数少,无导水主细胞分化,也没有厚壁细胞,这些特点使得水灰藓能够生长在水生环境中;东亚小金发藓Pogonatum inflexum和波叶仙鹤藓Atrich umundulatum的叶腹面覆盖着栉片,东亚砂藓Racomitrium japonicum、大羽藓Thuidium cym-bifolium、福氏蓑藓Macromitrium ferriei、东亚短颈藓Diphyscium fulvifolium、扭口藓Barbula unguiculata和角齿藓Ceratodon purpureus的叶片表面有乳头状突起或疣状物,这些附属结构使它们能够适应于旱生的环境中。     

Ectopic divisions in vascular and ground tissues of Arabidopsis thaliana result in distinct leaf venation defects

Leaf venation patterns vary considerably between species and between leaves within a species. A mechanism based on canalization of auxin transport has been suggested as the means by which plastic yet organized venation patterns are generated. This study assessed the plasticity of Arabidopsis thaliana leaf venation in response to ectopic ground or procambial cell divisions and auxin transport inhibition (ATI). Ectopic ground cell divisions resulted in vascular fragments between major veins, whereas ectopic procambial cell divisions resulted in additional, abnormal vessels along major veins, with more severely perturbed lines forming incomplete secondary and higher-order venation. These responses imply limited vascular plasticity in response to unscheduled cell divisions. Surprisingly, a combination of ectopic ground cell divisions and ATI resulted in massive vascular overgrowth. It is hypothesized that the vascular overproduction in auxin transport-inhibited wild-type leaves is limited by simultaneous differentiation of ground cells into mesophyll cells. Ectopic ground cell divisions may negate this effect by providing undifferentiated ground cells that respond to accumulated auxin by differentiation into vascular cells.     

Leaf and twig anatomy of Eriope, a xeromorphic genus of Labiatae

The leaf and twig anatomy of 25 species of the genus Eriope were studied. The twig anatomy is very uniform apart from the level of formation of early layers of cork. Leaf anatomy shows considerable variation between the species, and this is correlated to some extent with the extreme habit range from trees to woody herbs. Characters of the lamina that show variation are: trichome type and frequency, cuticular markings, leaf dorsiventral or isobilateral, presence of adaxial stomata, presence of a hypodermis, number of layers of adaxial palisade mesophyll cells, occurrence of large bundles of phloem fibres at main veins, type of areolation and marginal venation. Petiole vasculature is simple and generally with either four distinct vascular bundles or two vascular arcs. The most xeromorphic species are usually woody herbs or sub-shrubs, and tend to have thick, isobilateral leaves with large bundles of phloem fibres and few hairs, or strongly dorsiventral leaves with a hypodermis and stomata in deep abaxial hair-lined depressions. The correlation of xeromorphic characters with environmental conditions is discussed. Leaf anatomy is of limited value in elucidating relationships within the genus.     

Developmental process of sun and shade leaves in Chenopodium album L.

The authors’ previous study of Chenopodium album L. revealed that the light signal for anatomical differentiation of sun and shade leaves is sensed by mature leaves, not by developing leaves. They suggested that the two‐cell‐layered palisade tissue of the sun leaves would be formed without a change in the total palisade tissue cell number. To verify that suggestion, a detailed study was made of the developmental processes of the sun and shade leaves of C. album with respect to the division of palisade tissue cells (PCs) and the data was expressed against developmental time (leaf plastochron index, LPI). The total number of PCs per leaf did not differ between the sun and shade leaves throughout leaf development (from LPI ?1 to 10). In both sun and shade leaves, anticlinal cell division of PCs occurred most frequently from LPI ?1 to 2. In sun leaves, periclinal division of PCs occurred synchronously with anticlinal division. The constancy of the total number of PCs indicates that periclinal divisions occur at the expense of anticlinal divisions. These results support the above suggestion that two‐cell‐layered palisade tissue is formed by a change of cell division direction without a change in the total number of PCs. PCs would be able to recognize the polarity or axis that is perpendicular to the leaf plane and thereby change the direction of their cell divisions in response to the light signal from mature leaves.     

Clonal analysis of stomatal development and patterning in Arabidopsis leaves.

Cell lineage has been used to explain the stomatal distribution in several plant species. We have used transgenic plants carrying a 35SGUS::Ac construct that produces clonal sectors to analyze the possible role of cell lineage during the establishment of stomatal patterning in Arabidopsis leaves. The analysis of sectors ranging from two to eighteen cells supports the conclusion that most stomatal complexes derive from a single and immediate precursor cell through a stereotyped pattern of three unequal cell divisions followed by a final equal one. In addition, it shows that the successive cell divisions take place at a constant angle (approximately 60 degrees ) with respect to the previous one. Interestingly, this angular dimension shifts from 60 degrees to 0 degrees in the last cell division that gives rise to the stoma. These sectors also reveal the development of both clockwise and counterclockwise patterns of cell divisions during stomatal development in approximately equal numbers. Our clonal analysis indicates that cell divisions involved in the development of stomatal complexes are probably the last ones contributing to epidermal growth and development. Finally, the stereotyped pattern of cell divisions that culminates in the formation of stomatal complexes indicates that cell lineage plays a very important role during stomatal pattern establishment.     

DEVELOPMENTAL ANATOMY OF DIRECT SHOOT ORGANOGENESIS FROM LEAF PETIOLES OF VITIS VINIFERA (VITACEAE)

The anatomy of direct shoot organogenesis from leaf petioles of Vitis vinifera cv. French Colombard cultured in vitro was studied by light microscopy. Regenerating petiole stubs were fixed at 2- or 3-day intervals and sectioned longitudinally. By day 3 on regeneration medium, new cell divisions were observed. After 6 days, three distinct regions of meristematic activity were apparent within the expanding petiole stub: the wound-response, organogenic, and vascularization regions. In the organogenic region, rapid periclinal divisions of vacuolate outer cortical cells formed nodular bumps, many of which developed vascular strands and marginal meristems and formed adventitious leaves. Promeristems with small, densely staining cells and a distinct tunica layer also originated in the organogenic region, by cell division in the epidermal and subepidermal cell layers. With vascularization and the formation of leaf primordia, many promeristems became adventitious shoot meristems. Adventitious leaves and promeristems were initiated continuously from day 10 until day 33. Promeristems were often initiated near or upon adventitious leaves but could form either before or after the adventitious leaf developed. Adventitious leaves and shoot meristems developed vascular connections with the vascular bundles of the original expiant. The implication of this pattern of regeneration for Agrobacterium-mediated transformation of Vitis is discussed.     

Structure and development of the secretory cavities of Myrtus communis leaves

The structure and development of Myrtus communis L. secretory cavities has been studied in young and expanded leaves, using light and scanning electron microscope. Secretory cavities are continuously formed during leaf development, but in mature leaves the rhythm of their appearance shows steep decrease. Each secretory cavity is developed from a single epidermal cell, which undergoes a periclinal division followed by anticlinal and several oblique cell divisions. The lumen of the secretory cavity is initiated by cell wall separation, i.e., schizogenously. The secretory cells line the cavity, where the secreted material is collected. Secretory cavities are covered by modified epidermal cells, which do not seem to form any special aperture. Essential oils seem to be discharged after mechanical treatment of the leaf.     

四种藓类植物叶片解剖结构观察

使用石蜡切片法对四川壤塘的 4种藓类植物叶片的结构进行解剖学观察和比较分析 ,结果表明扭口红叶藓Bryoerythrophylluminaequalifolium (Tayl.)Zand .、台湾拟金发藓Polytrichas trumformosum (Hedw .)G .Sm .和梨蒴珠藓BartramiapomiformisHedw .的中肋由多层细胞构成 ,并有明显的细胞分化 ,而长蒴紫萼藓GrimmiamacrothecaMitt.的中肋结构较简单 ,无明显的细胞分化。 4种藓类植物的叶片细胞都只有 1层 ,但台湾拟金发藓的叶片上有栉片 ,扭口红叶藓叶片细胞表面有细疣 ,长蒴紫萼藓的叶片细胞壁明显加厚。此外 ,4种植物的叶片厚度和中肋厚度 ,及其细胞密度之间差异都比较明显     

Aerobryum brevicuspis S.He (Brachytheciaceae), a new species from northern Vietnam

A new species of Aerobryum Dozy & Molk. (Brachytheciaceae), A. brevicuspis S.He from Lao Cai Province, northern Vietnam is described and illustrated. The new species resembles epiphytic A. speciosum Dozy & Molk. in the presence of pendent, sparsely branched stems, similar shape of leaves and areolation, but differs by its apiculate to cuspidate leaf apices, a single, faint or sometimes double costae in branch leaves, rather differentiated alar cells, the presence of a central strand in the stems, and numerous clustered axillary hairs that are often 5-8(-10) cells long.     

On the benefits of living in clumps: a case study on Polytrichastrum formosum

• The study concerns the mechanics and water relationships of clumps of a species of endohydric moss, Polytrichastrum formosum.
• Anatomical and morphological studies were done using optical and scanning electron microscopy. Experiments on waterdrop capture and their distribution to adjacent shoots within a moss clump were performed with the experimental set‐up for the droplet collision phenomena and ultra‐high speed camera. The mechanical strength of the moss clump was tested on an electromechanical testing machine.
• During the process of moss clump wetting, the falling water drops were captured by the apical stem part or leaves, then flowed down while adhering to the gametophore and never lost their surface continuity. In places of contact with another leaf, the water drop stops there and joins the leaves, enabling their hydration. Mathematical analysis of anatomical images showed that moss stems have different zones with varying cell lumen and cell wall/cell radius ratios, suggesting the occurrence of a periodic component structure. Our study provides evidence that the reaction of mosses to mechanical forces depends on the size of the clump, and that small groups are clearly stronger than larger groups.
• The clump structure of mosses acts as a net for falling rain droplets. Clumps of Polytrichastrum having overlapping leaves, at the time of loading formed a structure similar to a lattice. The observed reaction of mosses to mechanical forces indicates that this phenomenon appears to be analogous to the ‘size effect on structural strength’ that is of great importance for various fields of engineering.

     

Leaf initiation and development in soybean under phosphorus stress

Experiments investigated changes in leaf development in young soybean plants progressing into P stress. The apical meristem and leaf structure were examined anatomically to evaluate the involvement of cell division and cell expansion in the restriction of leaf number and individual leaf size. Seedlings were deprived of P for 32 d following germination. Leaf initiation rates declined noticeably after about 2 weeks, even though the apical dome was of similar size and had a similar number of cells as controls. Primordia appeared morphologically similar also. Expansion of primary and the first three trifoliolate leaves of -P plants was severely reduced, and expansion of each leaf ceased, uniformly, when an area of about 40 cm(2) was obtained. Leaf epidermal cell size in the lateral plane was unaffected. The results indicate that expansion of leaves under P stress was limited by the number of cell divisions, which would imply control of cell division by a common regulatory factor within the leaf canopy.     

FLEXIBILITY IN ONTOGENY AS SHOWN BY THE CONTRIBUTION OF THE SHOOT APICAL LAYERS TO LEAVES OF PERICLINAL CHIMERAS

The development of leaves on apically stable, periclinal chimeras was studied in a number of dicot genera. The mutant cell layers of the shoot apex and the tissues derived from them were as active developmentally as the normal layers. Ontogeny was the same in these chimeras as in nonchimeras, and growth of their leaves can be outlined as follows. Formation of the buttress, the axis, and the lamina of simple dicot leaves were independent events. In each the first growth included derivatives of the apical layers, usually three in number, found in the apex of the shoot and the lateral buds. Most cell divisions in the outer layers (L-I and L-II) were anticlinal relative to the new structures. Therefore, in the proximal regions of the buttress, axis (petiole and midrib), and lamina, the derivative cells of L-I and L-II were usually present in single layers. The rest of the internal tissue was from L-III. As formation of the axis and the lamina proceeded, derivatives of L-II replaced L-III internally in the distal and marginal regions leaving cells of L-III behind. Both the determinate growth of leaves and the pattern of cell divisions at and near the leading edges of growth meant that no cells in the leaf were comparable to the initial cells of the shoot apex. As the lamina extended, there were extensive intercalary cell divisions, both anticlinal and periclinal, so that in any given region of a leaf the layers of internal cells were from either L-II or L-III. At any point along the axis, L-III participated or did not participate in laminar extension. At any given stage in laminar growth either of two sister cells in any internal layer divided either a few times or extensively. The extreme variability in direction and frequency of cell division during leaf development was under an overriding genetic control, which resulted in the normal or typical size, shape and thickness of leaves.     

DEVELOPMENT OF THE DIMORPHIC CHLOROPLASTS OF SUGAR CANE

The vascular bundle sheath cells of sugar cane contain starch-storing chloroplasts lacking grana, whereas the adjacent mesophyll cells contain chloroplasts which store very little starch and possess abundant grana. This study was undertaken to determine the ontogeny of these dimorphic chloroplasts. Proplastids in the two cell types in the meristematic region of light-grown leaves cannot be distinguished morphologically. Bundle sheath cell chloroplasts in tissue with 50% of its future chlorophyll possess grana consisting of 2-8 thylakoids/granum. Mesophyll cell chloroplasts of the same age have better developed grana and large, well structured prolamellar bodies. A few grana are still present in bundle sheath cell chloroplasts when the leaf tissue has 75% of its eventual chlorophyll, and prolamellar bodies are also found in mesophyll cell chloroplasts at this stage. The two cell layers in mature dark-grown leaves contain morphologically distinct etio-plasts. The response of these two plastids to light treatment also differs. Plastids in tissue treated with light for short periods exhibit protrusions resembling mitochondria. Plastids in bundle sheath cells of dark-grown leaves do not go through a grana-forming stage. It is concluded that the structure of the specialized chloroplasts in bundle sheath cells of sugar cane is a result of reduction, and that the development of chloroplast dimorphism is related in some way to leaf cell differentiation.     

LEAF DIMORPHISM IN POPULUS TRICHOCARPA

Critchfield , William B. (Pacific SW Forest & Range Expt. Sta., Berkeley, Calif.) Leaf dimorphism in Populus trichocarpa. Amer. Jour. Bot. 47 (8) : 699–711. Illus. 1960.—In Populus trichocarpa and other species of Populus, each tree bears 2 kinds of leaves, referred to here as “early” and “late” leaves. Both leaf types are present on all long shoots. They differ in many features of external morphology, including petiole length, size and occurrence of marginal glands, venation, and stomatal distribution. This type of foliar dimorphism has its origins in a pronounced difference in leaf ontogeny. The early leaves originate in the developing bud and overwinter as embryonic leaves. The first late leaves are also present in the winter bud, but as arrested primordia, and succeeding late leaves are initiated at the tip of the growing shoot and develop uninterruptedly to maturity during the growing season. A similar correlation between leaf form and the circumstances of leaf ontogeny appears to be a common feature of many other instances of heterophylly. The expansion of the pre-formed early leaves is almost completed by late spring, when the first late leaves begin to grow rapidly. The formation of late leaves may then continue until late in the season. The rapid elongation of the stem does not begin until the first late leaves expand. Elongation is restricted to shoots producing late leaves. Consequently, the early leaves are confined to short shoots and the base of long shoots; adventitious shoots and the upper part of long shoots bear only late leaves. Certain other woody plants with long and short shoots also exhibit a restriction of elongation to those shoots on which a second set of leaves is produced.     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

A Sequential Study of Cell Divisions and Expansion Patterns on a Single Developing Shoot Apex of Vinca major

During the growth of a single developing vegetative apex ofVinca major, both the orientation and frequency of cell divisions,and the pattern of cell expansion, were observed using a non-destructivereplica technique. Micrographs taken at daily intervals illustratethat the central region of the apical dome remains relativelyinactive, except for a phase of cell division which occurs after2 d of growth. The majority of growth takes place at the proximalregions of the dome from which develop the successive pairsof leaves. The developing leaf primordia are initiated by aseries of divisions which occur at the periphery of the centraldome and are oriented parallel to the axis of the subsequentleaves. The cells which develop into the outer leaf surfaceof the new leaves undergo expansion and these cells divide allowingfor the formation of the new leaf. This paper describes thefirst high-resolution sequential study of cell patterns in asingle developing plant apex. Sequential development, cell division, expansion patterns, SEM, Vinca major, apical dome, leaf primordium, leaf initiation     

Organ shape and size: a lesson from studies of leaf morphogenesis

Control of the shape and size of indeterminate organs, such as roots and stems, is directly related to the control of the shape and size of the cells in these organs, as predicted by orthodox cell theory. For example, the polarity-dependent growth of leaf cells directly affects the polar expansion of leaves. Thus, the control of leaf shape is related to the control of the shape of cells within the leaf, as suggested by cell theory. By contrast, in determinate organs, such as leaves, the number of cells does not necessarily reflect organ shape or size. Genetic evidence shows that a compensatory system(s) is involved in leaf morphogenesis, and that an increase in cell volume can be triggered by a decrease in cell number and vice versa. Studies of chimeric leaves also suggest interaction between leaf cells that coordinates the behaviour of these cells at the organ level. Moreover, leaf size also appears to be coordinated at the whole-plant level. The recently hypothesised neo cell theory describes how leaf shape- and size-control mechanisms control leaf shape at the organ-level via cell-cell interaction.