Independence of Organogenesis and Cell Pattern in Developing Angle Shoots of Selaginella martensii

Angle meristems are mounds of meristematic tissue located atdorsal and/or ventral branch points of the dichotomising stemaxes of many species of Selaginella (Lycophyta). The presentstudy examined the development of ventral angle shoots of S.martensii in response to removal of distal shoot apices (decapitation).Scanning electron microscopy of sequential replicas of developingangle meristems and angle shoots revealed that for the firsttwo pseudowhorls of leaf primordia, particular leaves are notattributable to particular merophytes of the angle meristemapical cell. Individual leaf primordia of the first (outer)pseudowhorl often form from more than one merophyte. Neitherthe shape of the angle meristem apical cell nor the directionof segmentation has any effect on the development of the angleshoot. Additionally, the apical cell of the angle meristem doesnot necessarily contribute directly to either of the new shootapices of the developing angle shoot. The first bifurcationof the angle shoot shows a remarkably consistent relationshipto the branching pattern of the parent shoot. The strong branchof the first angle shoot bifurcation typically occurs towardthe weak side branch of the parent shoot. Anatomical studiesshowed that bifurcation of the young angle shoot involved theformation of two new growth centres some distance away fromthe original angle meristem apical cell; new apical cells subsequentlyformed within these. These results provide additional supportfor the view that cell lineage has little or no effect on finalform or structure in plants.Copyright 1994, 1999 Academic Press Selaginella martensii Spring, Lycophyta, angle meristem, apical cell, shoot apical meristem, leaf primordium, branching, dichotomy, morphogenesis, determination, competence, development, mould and cast technique, replica technique, scanning electron microscopy     

Angle Meristem Origin and Development in Selaginella martensii

The origin and early development of angle meristems (AMs) inSelaginella martensii were studied using a high-resolution mouldand cast technique for scanning electron microscopy, in conjunctionwith light microscopy. Both dorsal and ventral AMs originatein the apical region shortly after the most recent bifurcationof the shoot apex. AMs appear to arise from a single superficialinitial cell, which subsequently undergoes an oblique anticlinaldivision to establish the apical cell of the AM. The AM apicalcell has a relatively short active life and divides laterallyusually only six to twelve times; most cells of the AM originatefrom repeated divisions of apical cell derivatives. Dorsal andventral AMs show similar developmental sequences, although theydiffer slightly in time of appearance. Ventral AMs originatedfirst, dorsal AMs appearing shortly thereafter. At a given branchpoint, the ventral AM is typically larger than the correspondingdorsal AM. Both dorsal and ventral AMs grow out as rhizophores(aerial roots), although ventral AM development greatly precedesdorsal AM development. It is possible that the apparent earlierdecrease in apical cell activity in dorsal AMs compared to ventralAMs could account for the lag in dorsal AM development as arhizophore at a given branch point. Selaginella martensii Spring, Lycophyta, angle meristem, apical cell, morphogenesis, mould and cast technique, replica technique, scanning electron microscopy     

LEAF DEVELOPMENT AND PRIMARY VASCULAR ORGANIZATION IN SHOOTS OF ANISOPHYLLEA DISTICH A

Anisophyllea disticha is characterized by strong shoot dimorphism. Orthotropic shoots with helically arranged scale leaves produce tiers of plagiotropic shoots, while plagiotropic shoots are anisophyllous and bear dorsal scale and ventral foliage leaves arranged in a unique tetrastichous system. In this study we compare the patterns of leaf development and primary vascular organization in the two types of shoots. Orthotropic shoots have an open vascular system with five sympodia. Expansion of orthotropic shoot scale leaves occurs from P1 to P10–12, and leaf tissues mature precociously. Plagiotropic shoots have a closed vascular system with six sympodia. Leaves in ventral and dorsal orthostichies do not differ significantly in size until ca. P15, but ventral leaves are distinct histologically from the second node in an orthostichy, P4–6. Ventral foliage leaves have a diffuse plate meristem, and leaf expansion continues until ca. P30. Differentiation of ventral and dorsal leaf trace procambium parallels the divergent patterns of leaf expansion. These observations demonstrate the strong correlation among shoot symmetry, leaf development, and vascular differentiation within dimorphic shoots of one species.     

EARLY LEAF ONTOGENY AND SHOOT DEVELOPMENT IN BROWNEA ARIZA (LEGUMINOSAE)

Brownea ariza Benth. (Leguminosae: Caesalpinioideae) shows early shoot tip abortion and subsequent renewal growth from the pseudoterminal bud. This species is unusual in that the entire shoot system is formed before flushing from the bud occurs, shoot tip abortion occurs during flushing, and the aborting portion contains three to six leaves as well as primordial structures varying from hood to peg shape. This study focused on the morphological changes from initiation of scale and foliage leaf primordia in the “resting” renewal bud through bud elongation to flushing and bud abortion. Scanning electron microscopy revealed that embryonic scale leaves are hood-shaped while foliage leaf primordia show early segmentation into leaflets and stipules. No transitional stages were observed. Bud scales and foliage leaves show opposite developmental trends. In bud scales, length at maturity increases from first to last formed, while length decreases in sequentially formed foliage leaves. Early in leaf development the stipules keep pace with the elongation of the rachis. When the bud reaches about one half of its final length the leaf rachis begins to exceed the lengths of its stipules. This young rachis terminates in a distinct mucro that persists until maturity at which time it abscises. Growth patterns indicate that mucro and rachis are a single developmental unit. The early abortion of a shoot tip containing several leaves cannot be easily rationalized. Previous suggestions have involved maintenance of form and ecological adaptation. We add the possibility of elimination of cell progeny encumbered by mutations. From this and other studies of this group, it is clear that at maturity leaves of different species may look alike, e.g., Hymenaea and Colophospermum are bifoliolate; Brownea, Saraca, and others are multifoliolate. However, early stages of leaf ontogeny are quite diverse and may be of systematic value, since these early differences are lost or masked by later development.     

LEAF DEVELOPMENT IN ISOPHYLLOUS AND FACULTATIVELY ANISOPHYLLOUS SPECIES OF PENTADENIA (GESNERIACEAE)

Shoots of Pentadenia orientandina exhibit varying degrees of anisophylly, ranging from pairs of equal-sized leaves to pairs of large ventral and small dorsal leaves. In this study we compare phyllotaxis, leaf expansion, and accompanying histological changes in extremely anisophyllous shoots of this species and in isophyllous shoots of the related species, P. crassicaulis. In P. orientandina, decussate phyllotaxis is modified at leaf initiation, and angles of leaf insertion appear to be further changed during leaf expansion. In both species, leaf primordia of a pair are not distinguishable at inception, suggesting an equivalent developmental potential. In P. orientandina, size differences between ventral and dorsal leaves become significant at the P2 or P3 stage, coincident with lamina initiation. Minute dorsal leaves are arrested in their development at the P3 stage and mature without differentiation of multiple epidermis, stomata, mesophyll and most vascular tissue. Variation in dorsal leaf structure in P. orientandina emphasizes the plasticity of leaf development in this facultatively anisophyllous species.     

LEAF DIMORPHISM IN THE AQUATIC ANGIOSPERM CALLITRICHE HETEROPHYLLA

Depending on the position of the shoot tip relative to the water surface, the aquatic angiosperm Callitriche heterophylla produces either ovate land-form or linear water-form leaves. This paper is concerned with the developmental basis for the leaf dimorphism of this species. Little significant difference is observed between the apical meristems of submerged vs. emergent shoots; moreover, land-form and water-form primordia undergo similar, if not identical, patterns of initial development until they attain a length of 350 to 400 μm. These findings are interpreted to mean that the divergent leaf forms result from the marked sensitivity of the primordia to their respective environments rather than from the mode of their inception. Subsequent growth of the young water-form leaf emphasizes longitudinal extension, while the immature land-form leaf continues balanced expansion in both longitudinal and lateral directions. The lateral growth of the land-form primordium is accomplished in part by a more persistent marginal meristem, but the morphological difference between the two leaf forms is mostly attributable to the difference in the predominant direction of intercalary expansion. In addition, certain anatomical features, such as vasculature, stomates, and cuticle, are much more prominent in mature land-form leaves than in water-form leaves. These anatomical differences seem to represent structural adaptations of each leaf form to the specific physiological requirements of its environment.     

A MORPHOMETRIC STUDY OF THE ULTRASTRUCTURE OF ECHINOCEREUS ENGELMANNII (CACTACEAE). IV. LEAF AND SPINE PRIMORDIA

A stereological morphometric study of leaf primordia (P1 and P2) of Echinocereus engelmannii indicated that primordia are significantly different ultrastructurally from the shoot apical meristem tissues (tunica and peripheral zone) that produce the primordia. Leaf initiation involves readjustments of rates of synthesis and growth of cytoplasm, vacuoles, mitochondria, chloroplasts, and dictyosomes, such that leaf initiation must be a complex process in which different cell components are affected individually. Furthermore, leaf primordia are ultrastructurally distinct from spine primordia. Leaf and spine primordia as young as these are not yet irrevocably determined, thus different types of primordia, from the time of their inception and before their determination, have distinctly unique metabolisms; primordia are not merely generalized, uncommitted outgrowths whose developmental fate is set at some time later than inception.     

Growth of Populus euramericana

Growth curves of successive leaves of Populus euramericana (Dode) Guinier 'Robusta' have been determined. With ample supply of water and nutrients the growth of a poplar shoot follows a fixed pattern: an initial logarithmic acceleration phase followed by a stationary phase in which leaves of equal size are produced at a constant rate. Analysis of growth curves of leaves enabled the growth curves of leaf primordia to be predicted. These primordial growth curves are compared to the indirectly determined growth curves of primordia by measuring the lengths of successive leaf primordia in the apex during the stationary phase of growth. The increase in length of successive leaves in the acceleration phase of growth continues for a longer period at high than at low irradiace. The relative growth rates of leaf primordia, leaves and internodes are discussed in terms of shoot growth and phyllotaxis.     

DEVELOPMENTAL ANALYSIS OF LEAF PLASTICITY IN THE HETEROPHYLLOUS AQUATIC PLANT HIPPURIS VULGARIS

Leaf development was studied in the heterophyllous aquatic plant Hippuris vulgaris in order to characterize the developmental events that lead to the formation of aerial- vs. submerged-type leaves. Recent evidence that abscisic acid regulates leaf development in this species provided a basis for using abscisic acid as a developmental tool to accurately control leaf development. We found that leaf primordia were fully competent to develop into either aerial- or submerged-type leaves until the 10th plastochron, when they were ca. 300 μm long. Also, leaves between about the 10th and 21st plastochron formed sectored transition leaves (i.e., the basipetal portion was composed of aerial-type tissue and the apical portion was composed of submerged-type tissue, or vice versa), indicating that tissue determination as one or the other leaf type occurred on a local, as opposed to whole-leaf, level. Finally, no significant difference was observed between the apical dimensions of aerial or submerged-type shoots. These results indicate that the final determination of Hippuris vulgaris leaves occurs a) relatively late in leaf development, and b) independently of the shoot apex, and provide a basis for using this plant in further studies concerning leaf determination and pattern formation (e.g., stomates, lateral venation) in plants.     

越南莲座状复苏卷柏一新种——越南卷柏(卷柏科)

该文描述了在越南发现的卷柏科一新种——越南卷柏(Selaginella pseudotamariscina X. C. Zhang & C. W. Chen)。经过分子系统发育分析并与近缘种形态比较,认为该新种是卷柏属(Selaginella)同穗亚属(subg. Stachygynandrum)的一个物种,与S. digitata-S. imbricata分支为姐妹群关系。该新种与卷柏(S. tamariscina)和垫状卷柏(S. pulvinata)的形态近似,其植株均为莲座状,但不同之处在于中叶对称,似披针形,上表面具1沟槽; 孢子叶穗略压扁; 孢子叶和营养叶性状和排列近似,孢子叶非同形,略异形,正置,腹面孢子叶大于背面孢子叶; 背面孢子叶败育,孢子囊仅见于腹面孢子叶基部。     

PHOTOTROPISM OF SELAGINELLA: THE DIFFERENTIAL RESPONSE TO LIGHT

Selaginella kraussiana (Kunze) A. Braun has a plagiotropic dorsiventral stem with two rows of small leaves on the dorsal surface and two rows of large leaves inserted laterally. Stem tips exhibit a differential phototropic response. When stem tips are placed in their normal horizontal orientation and the dorsal surfaces are illumianted, the tips bend only 20” below the horizon and away from light. Stem curvature is limited to a zone 450 μm long located 1.5 mm behind the shoot apex. The dorsal cortical cells within this zone of curvature are about 1.44 times longer than the ventral cortical cells. Illumination of the ventral surface of the stem tips elicits a strong phototropic response. The stems bend from 123–158° below the horizon and toward light, and the zone of curvature increases in length to 10 mm of the explant. The curvature is large enough so that the previously shaded dorsal leaves of the stem tips become redirected toward the light. This phototropic response is promoted by white and blue light, whereas red or far-red light has no effect. When stem tips are cultured in total darkness, the length of the zone of curvature is 8.0 mm but the stems bend only 50–67°. Treatment of the small dorsal leaves with phenylacetic acid inhibits phototropic curvature, and the phototropic response is unaffected by gravity.     

VASCULAR ARCHITECTURE IN ISOPHYLLOUS AND FACULTATIVELY ANISOPHYLLOUS SPECIES OF PENTADENIA (GESNERIACEAE)

The organization and differentiation of primary vascular tissue in isophyllous shoots of Pentadenia crassicaulis and facultatively anisophyllous shoots of P. orientandina (Gesneriaceae) were compared using serial reconstructions and quantitative methods. Despite clear differences in shoot symmetry, both species are vascularized by four sympodia, with trilacunar, split-lateral nodal anatomy. Leaf trace tracheary element number and diameter reflect leaf size differences in P. orientandina: these parameters are significantly greater in the large ventral leaves than in the small dorsal leaves. The median and lateral traces of ventral leaves of this species have a similar number of tracheary elements of equal diameter, while there are significantly more tracheary elements in the median than lateral traces of dorsal leaves. The pattern seen in P. crassicaulis is similar to that seen in the dorsal leaves of P. orientandina. In both species, protoxylem development anticipates differences in mature shoot vasculature. Changes in tracheary element number during ontogeny precede or are approximately coincident with changes in leaf size. These results suggest that the facultative expression of leaf size differences in P. orientandina is associated with opportunistic development and differentiation of the lateral trace.     

ORGANOGRAPHY,BRANCHING, AND THE PROBLEM OF LEAF VERSUS BUD DIFFERENTIATION IN THE VINING EPIPHYTIC FERN GENUS MICROGRAMMA

Investigation of the development and organography of the shoot systems of Microgramma vacciniifolia and M. squamulosa was undertaken for the purpose of determining: (1) the features of shoot growth that are responsible for the distinctive vining character of these epiphytic ferns; and (2) the mode of origin of branches and their contrast with leaf initiation. Shoots of both species are dorsiventral and plagiotropic (i.e., parallel to the substrate) in habit. Since the shoot apical meristem is radial in transectional symmetry, shoot dorsiventrality in Microgramma is a postgenital or secondary developmental event, and its inception is related to the initiation of lateral appendages. Leaves and buds arise in a distichous phyllotaxis and occupy opposite and alternating positions on the dorsal surfaces and flanks of the rhizome. Endogenous roots are initiated in two rows from the ventral surface of the stem, in the vicinity of the rhizome meristem; however, they do not emerge from the rhizome until some distance behind the tip and do not elongate until the region of substrate contact. We conclude that the vining nature of this fern rhizome is a result of precocious internodal elongation and the concomitant delay of leaf and bud expansion in the region of stem elongation. In addition, observation of branch origin confirms previous suggestions that branching in Microgramma is strictly lateral and extra-axillary and not a dichotomous derivative as proposed by some workers. Leaf and bud primordia differ not only in the nature of their respective vascular supplies but also in their actual course of initiation. In the case of the leaf, the primordium is precociously emergent and exhibits a lenticular apical cell at its summit when it is only one plastochron removed from the flanks of the apical meristem. By contrast, initials of the bud primordium divide less actively and remain in a sunken position for at least 5–6 plastochrons; only when the bud apex becomes expanded and emergent does a tetrahedral apical cell become recognizable at the tip of the bud promeristem. Because of the distinctive pattern of branch and leaf origin, as well as the lack of adventitious and phyllogenous origin of branch primordia, we suggest that the shoot of Microgramma is a useful test organism for the re-examination of the problem of leaf and bud determination in the ferns.     

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.     

Effects of irradiation level on leaf growth of sunflower

Sunflower, Helianthus annuus L. cv. INRA 6501, plants were grown in a gravel culture subirrigated with Hoagland nutrient solution, at photosynthetically active radiation levels of 15, 30 and 60 W m^-2 at a daylength of 16 h, a temperature of 20°C and a relative humidity of 60% throughout. Development of the plant and growth of the leaves were measured. High irradiance accelerated development proportionally in all phases from germination, through leaf initiation, primordial flower formation and the maturation of all plant organs until anthesis. High irradiance levels stimulated the expansion of the growing shoot, which produced more and larger primordia. Under constant conditions the ratio between leaf initiation rate and mature length of a leaf remained constant, although the growth patterns [relationship between relative growth rate (RGR) and organ age] of successive leaves were not similar. Consequently, it may be assumed that, as in poplar, the increasing size of the growing shoot reflects the increase of the vascular system of sunflower. The growth patterns of the leaves depend on the developmental stage of the plant and, in the young primordial stage, also on irradiance level. In the linear phase of growth the growth pattern is independent of irradiance level.     

THE SHOOT-GROWTH RHYTHM OF A TROPICAL TREE,THEOBROMA CACAO

The shoot-growth rhythm of tropical trees is a little understood phenomenon. Correlations of tree growth with environment, determined from field studies in the tropics, have been largely inconclusive, and few studies have been done under controlled environmental conditions. As an initial part of a project to study shoot-growth rhythms in tropical trees this paper describes the rhythm in Theobroma cacao L. An individual shoot passes through alternate periods of growth and dormancy. The growth period is characterized by the expansion of leaves and elongation of the shoot. During dormancy the length of the shoot remains constant, and no new leaves expand. Shoot-growth rhythm was divided into phases. Dissection of shoot tips from the various phases shows that the total number of leaves and leaf primordia in the shoot apex remains constant during the dormant period and does not increase until the onset of the growth period. This indicates that activity of the apical meristem as well as leaf expansion and shoot elongation are rhythmic. We found that the rhythm of shoot growth persists under controlled environmental conditions and that growth under these conditions is asynchronous, as it appears to be in the field. Our data strongly suggest endogeneity.     

THE ROLE OF LIGHT IN HISTOGENESIS AND DIFFERENTIATION IN THE SHOOT OF PISUM SATIVUM. II. THE LEAF

Thomson , Betty F., and Pauline Monz Miller . (Connecticut Coll., New London.) The role of light in histogenesis and differentiation in the shoot of Pisum sativum, II. The leaf. Amer. Jour. Bot. 49(4): 383–387. Illus. 1962.—Development of the form and anatomy of leaves was studied in plants of Pisum sativum grown in vermiculite under constant conditions and exposed daily to red or white light or kept in continuous darkness. The red light used had an intensity in the morphogenetically active red region of the spectrum of 70–75% that of the white light. Light had no effect on the manner of initiation or early development of leaf primordia. Quantitative data from older leaves showed that light has no effect on the pattern of later development but does affect the rate and extent of development. Under all light conditions, the length of the leaflet is closely correlated with the state of its internal anatomy. “Mature” etiolated leaves duplicate young stages of light-grown leaves. Mature leaves grown in red light duplicate not-quite-mature leaves grown in white light. The difference between white-light and red-light leaves is attributed here to light intensity and resembles that between sun and shade leaves.     

COMPARATIVE FOLIAR HISTOGENESIS IN ACORUS CALAMUS AND ITS BEARING ON THE PHYLLODE THEORY OF MONOCOTYLEDONOUS LEAVES

A comparative histogenetic investigation of the unifacial foliage leaves of Acorus calamus L. (Araceae; Pothoideae) was initiated for the purposes of: (1) re-evaluating the previous sympodial interpretation of unifacial leaf development; (2) comparing the mode of histogenesis with that of the phyllode of Acacia in a re-examination of the phyllode theory of monocotyledonous leaves; and (3) specifying the histogenetic mechanisms responsible for morphological divergence of the leaf of Acorus from dorsiventral leaves of other Araceae. Leaves in Acorus are initiated in an orthodistichous phyllotaxis from alternate positions on the bilaterally symmetrical apical meristem. During each plastochron the shoot apex proceeds through a regular rhythm of expansion and reduction related to leaf and axillary meristem initiation and regeneration. The shoot apex has a three- to four-layered tunica and subjacent corpus with a distinctive cytohistological zonation evident to varying degrees during all phases of the plastochron. Leaf initiation is by periclinal division in the second through fourth layers of the meristem. Following inception early growth of the leaf primordium is erect, involving apical and intercalary growth in length as well as marginal growth in circumference in the sheathing leaf base. Early maturation of the leaf apex into an attenuated tip marks the end of apical growth, and subsequent growth in length is largely basal and intercalary. Marked radial growth is evident early in development and initially is mediated by a very active adaxial meristem; the median flattening of this leaf is related to accentuated activity of this meristematic zone. Differentiation of the secondary midrib begins along the center of the leaf axis and proceeds in an acropetal direction. Correlated with this centralized zone of tissue specialization is the first appearance of procambium in the center of the leaf axis. Subsequent radial expansion of the flattened upper leaf zone is bidirectional, proceeding by intercalary meristematic activity at both sides of the central midrib. Procambial differentiation is continuous and acropetal, and provascular strands are initiated in pairs in both sides of the primordium from derivatives of intercalary meristems in the abaxial and adaxial wings of the leaf. Comparative investigation of foliar histogenesis in different populations of Acorus from Wisconsin and Iowa reveals different degrees of apical and adaxial meristematic activity in primordia of these two collections: leaves with marked adaxial growth exhibit delayed and reduced expression of apical growth, whereas primordia with marked apical growth show, correspondingly, reduced adaxial meristematic activity at equivalent stages of development. Such variations in leaf histogenesis are correlated with marked differences in adult leaf anatomy in the respective populations and explain the reasons for the sympodial interpretation of leaf morphogenesis in Acorus and unifacial organs of other genera by previous investigators. It is concluded that leaf development in Acorus resembles that of the Acacia phyllode, thereby confirming from a developmental viewpoint the homology of these organs. Comparison of development with leaves of other Araceae indicates that the modified form of the leaf of Acorus originates through the accentuation of adaxial and abaxial meristematic activity which is expressed only slightly in the more conventional dorsiventral leaf types in the family.     

HETEROPHYLLY AND NEOFORMATION OF LEAVES IN SUGAR MAPLE (ACER SACCHARUM)

Variation in leaf form and timing of leaf initiation were investigated in vigorous leader shoots of open-grown saplings and larger forest trees of sugar maple (Acer saccharum Marsh.). Winter buds of leader shoots usually contained 6 or 8 leaf primordia and embryonic leaves, whereas 12 to 18 leaves typically expanded along the shoots each year. Preformed (early) leaves differ in form from neoformed (late) leaves. As in some other Acer species, the first-formed late leaves have large angles of secondary lobe divergence and deeply indented sinuses. This pattern of heterophylly contributes to the multilayered nature of open-grown saplings and leader shoots of forest trees of sugar maple.