Procambial Strands Research Articles

ONTOGENY OF MAJOR VEINS IN THE LAMINA OF POPULUS DELTOIDES BARTR

The ontogeny of the major venation in the lamina of Populus deltoides Bartr. leaves was investigated in relation to the development of original procambial bundles, subsidiary bundles, and their derivatives. Serial sections and clearings were used to show that the midrib region is a composite structure consisting of several independent vascular bundles, each of which eventually diverges into the lamina to become a secondary vein. The sequence of events in the ontogeny of major secondary veins is: (1) an original procambial strand develops acropetally and becomes the precursor of the first vascular bundle of the midrib region of the lamina, (2) ground tissue at the forefront of acropetally developing subsidiary procambial bundles differentiates in a wavelike continuum; meristematic regions precede the acropetally developing procambial bundles, (3) discrete subsidiary bundles differentiate in the meristematic regions as they advance acropetally, (4) subsidiary bundles diverge obliquely in the lamina margin giving rise to the secondary veins in a basipetal fashion, and (5) subsequent differentiation and maturation of the secondary veins occurs within the lamina. The original procambial bundles and first‐formed subsidiary bundles become the secondary veins of the uppermost portions of the lamina, the next‐formed subsidiary bundles become the secondary veins of the middle portions of the lamina, and the last‐formed subsidiary bundles become the secondary veins of the lowermost portion of the lamina.

Floral development of Aponogeton natans and A. undulatus

The primordia of the floral appendages are initiated in an acropetal succession. Members of the same whorl appear nearly simultaneously. The gynoecial whorl and the two staminal whorls are trimerous, whereas the perianth consists only of two anteriolateral tepals. However, the posterior (adaxial) tepal may be present as an extremely reduced buttress whose growth becomes arrested immediately after its inception. If this somewhat questionable tepal rudiment is included we have a perfectly trimerous and tetracyclic flower with alternation of successive whorls. Subtending bracts of the flowers are completely missing in all developmental stages. While the tepal primordia are dorsiventral from their inception, the stamen and pistil (carpel) primordia originate as hemispherical mounds which become dorsiventral in subsequent stages of development. Each pistil (carpel) primordium becomes horseshoe shaped. As the margins grow up and contact they fuse postgenitally. No cross zone is formed. Placentation is submarginal. In A. natans eight ovules are formed and in A. undulatus only two arise; all ovules are bitegmic. The floral apices have a two-layered tunica up to the stage of pistil formation. The inception of all floral appendages (including the ovules) occurs by periclinal cell division in the second tunica layer. The third layer (corpus) may contribute to the formation of the stamens and pistils. Each appendage primordium receives only one procambial strand which begins to differentiate after the inception of the primordium. The questionable rudimentary tepal buttress lacks a procambial strand. Apparently it does not reach the developmental stage at which procambial induction occurs. From the point of view of floral development, the two species of Aponogeton differ drastically from members of the Alismatales studied so far. Among the Helobiae, the Aponogetonaceae appear to be most closely related to the Scheuchzeriaceae and the Juncaginaceae (Triglochinaceae).

Phyllotactic transitions in the vascular system of Populus deltoides bartr. as determined by 14C labeling

Populus deltoides seedlings progress through 2/5, 3/8, and 5/13 orders of phyllotaxis in attaining Plastochron Index 16 (PI 16). The manner in which the vascular system was reoriented during these phyllotactic transitions was determined by anatomical analysis of serial microsections, whereas the positions of the transitions were determined by (14)C labeling. The midvein at the tip of leaves representing plants of different PI and leaves of different Leaf Plastochron Index (LPI) was fed (14)CO2 photosynthetically, and primordia LPI 0 through LPI-9 were dissected from the buds and analyzed for (14)C. By combining the labeling data with the anatomical observations it was possible to reconstruct the vascular system of a plant of PI 16 and to locate the phyllotactic transitions. Both the anatomical and the labeling data showed a high degree of reproducibility among plants suggesting that the phyllotactic pattern to which the vascular system conforms may be programmed in the plant and transmitted acropetally through the developing leaves and procambial strands. The origin of new primordia and the concepts of orthostichy, ontogenetic helix, and Fibonacci sequence are discussed as they apply to the vascular system of P. deltoides.

Ovule and seed development in Pinus radiata: postmeiotic development, fertilization, and embryogeny

Development of ovule tissues in Pinus radiata after meiosis, fertilization, and embryogeny is comparable with that of other pines, but P. radiata takes longer to develop. Fertilization occurs 15 months after pollination and morphological embryo maturity is reached 5 months later. In ovules harvested in spring after meiosis, a curved band of small cells with dense cytoplasm extends from the chalazal end of the ovule to the vascular tissue of the ovuliferous scale. It is interpreted as a procambial strand, which in the next year, differentiates basipetally into elongated, thick-walled cells with degenerated nuclei.

Leaf development in Begonia hispida var. cucullifera with special reference to vascular organization

The leaf is initiated by periclinal divisions in the second cell layer of the apex. Early in development, it assumes an asymmetrical shape as a result of the non-median position of the apical meristematic region that forms the leaf tip. The other main lobes of the leaf are formed by two additional regions of growth to the side of the tip. In the second plastochron, primordia are initiated on the upper surface of the main ribs of the leaf and these often develop into large, leaf-like appendages. During the third plastochron, the leaf becomes covered by a dense layer of trichomes. Vascular development is well correlated with regions of growth. The median procambial strand extends diagonally into the skewed apical growth region and another strand branches from this and develops on the other side. Lateral procambial strands usually form in an abaxial to adaxial sequence in the petiole: one strand of the first pair to develop forms part of the median rib and the other forms a third main rib on the opposite side. After the initiation of primordia on the leaf surface, a 'ventral system' of procambia is developed supplying mainly the ventral rib surface and consequently the epiphyllous appendages. This consists of branches from the adaxial-most peripheral bundles in the petiole as well as a bundle in the centre. The vascular anatomy of the stem and nodal region is also described in relation to that of the leaf.

Developmental anatomy of foliage leaves, bracts, calyx and corolla inPharbitis nil

The structure and ontogeny of the foliage leaves, bracts, bracteoles, calyx and corolla ofPharbitis nil were investigated, with special reference to the development of the lamina and the procambium. Reproductive organs used are those of a terminal inflorescence and axillary flowers induced by a single 16 hr dark period given to the seedling. The foliage leaf consists of the petiole and the broad lamina. Bracts show various forms and structures, which fluctuate from a lower leafy bract to an upper scaly one in a terminal inflorescence. The sepal is scaly. The corolla is funnel-shaped, and composed of five wedge-shaped petals.In the lamina of the foliage leaf primordium, marginal growth is followed by active growth by the plate meristem, and procambial strands of lateral veins differentiate from the residual meristem. The primordium of the lowest bract of the terminal inflorescence has already been initiated before the dark period, and develops into the bract, the residual meristem disappearing after the treatment. The leafy bract shows marginal growth and growth by the plate meristem similar to that of the foliage leaf, but of short duration. The activity of marginal growth of the scaly bract and the sepal decreases rapidly and procambial strands of lateral veins differentiate acropetally from highly vacuolated cells. The activity of marginal growth of the petal decreases gradually, and derivatives of the marginal meristem divide as a plate meristem. The corolla tube is initiated by co-operation of interprimordial growth and marginal growth of petal primordia.

Floral development of Zannichellia palustris

What, at maturity, appears to be a bisexual flower in the axil of one of two subopposite leaves, is revealed as a fertile nodal complex with quite different organization. Three appendages develop at each nodal complex. The first girdles the stem and becomes at maturity a membranous sheath about the entire node. The second subtends the axillary meristem, which terminates as the staminate flower, and branches laterally as a renewal growth in the axil of a sterile appendage just below the stamen. The third appendage is subopposite the terminal meristem, which gives rise to the pistillate floral bud towards the staminate flower, and a renewal growth apex towards the appendage. This renewal growth apex repeats the entire pattern at almost a 90° shift to the right or left, depending on the shoot. The single stamen of the staminate flower develops as those studied in Potamogeton and Ruppia. The pistillate flower develops two carpel primordia, which become peltate before initiating a single ovule primordium on the adaxial portion (Querzone). The membranous envelope which covers the carpels at maturity is initiated at ovule inception, below one of the carpels. A peltate stigma differentiates on a short style and at maturity becomes broad and lobed. The renewal growth apex has a one-layered tunica. The membranous sheaths of the node and of the pistillate flower are primarily protodermal in origin, while the rest of the sterile and reproductive appendages arise through activity in subprotodermal cells. Procambial development is acropetal closely following primordial inception. Each organ (sterile or fertile) receives one procambial strand, except for the membranous sheath about the node and the one about the pistillate flower.

DEVELOPMENTAL STUDIES IN PTYCHOSPERMA (PALMAE). II. THE STAMINATE AND PISTILLATE FLOWERS

The staminate flower of Ptychosperma mooreanum and P. gracile has three imbricate sepals, three valvate petals, an androecium of 9–13 or 27–32 stamens, and a lageniform pistillode. Pistillate flowers have three imbricate sepals, but the three petals are also imbricate, and the three (‐seven) staminodes are irregular and sometimes basally fused. The ovoid pseudomonomerous pistil has a single locule with a pendulous ovule and three small distal stigmatic lobes. All floral organs are initiated by periclinal divisions in the second tunica layer of the floral apex, with divisions in the outer layer of the cortex also apparent in the initiation of stamens and staminodes. Primordia of floral organs differ in shape but can be related to those of other palm flowers. Sepals are initiated and develop successively, and during floral organogenesis are horizontally oriented covering other organs. Petals are also initiated successively, but rapidly attain equal size. Petals of staminate and pistillate flowers are initially similar, but subsequent prolonged activity of marginal meristems results in widely imbricate petals in pistillate flowers. The polyandric androecia are clearly trimerous. As stamen initiation commences, the apex is divided into sectors or primary primordia consisting of a narrow ridge opposite each sepal and a large triangular bulge opposite each petal. Stamens arise as secondary primordia in alternating antesepalous and antepetalous cycles. Variation in number of stamens is due primarily to variation in the number of primordia that arise in the wider lower antepetalous loci. The primordium of the pistillode is girdling but becomes trilobed later. The gynoecium is initiated as three unequal crescentic carpel primordia that are united by zonal growth early in development. A procambial strand develops acropetally into each floral organ, and additional strands are formed as each organ increases in size. Floral organs are histologically specialized. The appearance of tannins, raphides, and silica and the maturation of sclerenchyma are basipetal within each floral organ.

Developmental Studies in Ptychosperma (Palmae). I. The Inflorescence and the Flower Cluster

This paper describes inflorescence structure, including organogenesis of the panicle and flower clusters and vasculature of flowering branches, for two species of Ptychosperma, a genus of arecoid palms. The inflorescence is an infrafoliar panicle with up to four orders of branches in a spirodistichous arrangement conforming to an irregular one-half phyllotaxy. The primordium of the inflorescence is crescentic and the apex has two tunica layers, a group of central cells, and a rib meristem. The distal flower-bearing parts or rachillae of all branches develop acropetally early in ontogeny and are vertically oriented in the bud. Although these rachillae terminate branches of different sizes and orders, they are similar in size and in number of flower clusters produced. Internodes and lower parts of branches develop later. Bracts of four types are produced: a prophyll and empty peduncular bract, bracts which subtend lateral branches, bracts subtending triads, and floral bracteoles. The prophyll and peduncular bracts are tubular and completely closed around all branches until about three months before the flowers reach anthesis. Bracts subtending lateral branches and those that subtend triads enlarge by small amounts of apical, adaxial, and marginal growth to cover subtended apices during early ontogeny, but are small to absent at maturity. Flower clusters are triads of two lateral staminate and a central pistillate flower. Organogenesis indicates that the triad is a sympodial unit. Flowers develop successively, each floral apex bearing a bracteole that subtends the next flower. The vasculature of the inflorescence may be divided into two systems. Bundles of the main axis extend acropetally into the vertically oriented branches as they are initiated and form a central cylinder of larger bundles in each branch. Flower clusters are supplied by a peripheral system of smaller bundles that develop later in relation to the developing floral organs. Bundles of the peripheral system branch frequently, but branching levels are irregular. The irregular branching of peripheral bundles appears related to the phyllotaxy of the flower clusters and the random right or left position of the first flower of the triad. The level of branching of a bundle may depend on the position of a floral primordium with respect to an existing procambial strand. Three (-4) bundles supply each staminate flower and six (-10) the pistillate flower. The histologically specialized inflorescence has stomata and contains abundant starch. Tannins and raphides, spherical silica bodies, and various forms of sclerenchyma appear in sequence and apparently provide support and protection during the long exposure of the branches.

DEVELOPMENTAL STUDIES IN PTYCHOSPERMA (PALMAE). I. THE INFLORESCENCE AND THE FLOWER CLUSTER

This paper describes inflorescence structure, including organogenesis of the panicle and flower clusters and vasculature of flowering branches, for two species of Ptychosperma, a genus of arecoid palms. The inflorescence is an infrafoliar panicle with up to four orders of branches in a spirodistichous arrangement conforming to an irregular one‐half phyllotaxy. The primordium of the inflorescence is crescentic and the apex has two tunica layers, a group of central cells, and a rib meristem. The distal flower‐bearing parts or rachillae of all branches develop acropetally early in ontogeny and are vertically oriented in the bud. Although these rachillae terminate branches of different sizes and orders, they are similar in size and in number of flower clusters produced. Internodes and lower parts of branches develop later. Bracts of four types are produced: a prophyll and empty peduncular bract, bracts which subtend lateral branches, bracts subtending triads, and floral bracteoles. The prophyll and peduncular bracts are tubular and completely closed around all branches until about three months before the flowers reach anthesis. Bracts subtending lateral branches and those that subtend triads enlarge by small amounts of apical, adaxial, and marginal growth to cover subtended apices during early ontogeny, but are small to absent at maturity. Flower clusters are triads of two lateral staminate and a central pistillate flower. Organogenesis indicates that the triad is a sympodial unit. Flowers develop successively, each floral apex bearing a bracteole that subtends the next flower. The vasculature of the inflorescence may be divided into two systems. Bundles of the main axis extend acropetally into the vertically oriented branches as they are initiated and form a central cylinder of larger bundles in each branch. Flower clusters are supplied by a peripheral system of smaller bundles that develop later in relation to the developing floral organs. Bundles of the peripheral system branch frequently, but branching levels are irregular. The irregular branching of peripheral bundles appears related to the phyllotaxy of the flower clusters and the random right or left position of the first flower of the triad. The level of branching of a bundle may depend on the position of a floral primordium with respect to an existing procambial strand. Three (‐4) bundles supply each staminate flower and six (‐10) the pistillate flower. The histologically specialized inflorescence has stomata and contains abundant starch. Tannins and raphides, spherical silica bodies, and various forms of sclerenchyma appear in sequence and apparently provide support and protection during the long exposure of the branches.

DEVELOPMENT AND ORGANIZATION OF THE PRIMARY VASCULAR SYSTEM IN POPULUS DELTOIDES ACCORDING TO PHYLLOTAXY

An actively growing cottonwood bud was embedded in epon‐araldite and serially sectioned at 2 μm. The sections were analyzed microscopically with the optical shuttle system of Zimmermann and Tomlinson, and all data were quantitatively recorded relative to the apex and to leaf plastochron index (LPI). Analysis of the sections revealed an acropetally developing procambial system organized according to a precise phyllotaxy. Six procambial strands could be recognized and followed long before the leaf primordia that they would enter were evident at the apex. Origin of these strands coincided with developmental events both in the parent trace and its primordium and in the antecedent leaf on the same orthostichy. Once a primordium and its trace attained a certain stage of development, trace bundles began to develop basipetally from the primordium base. These trace bundles appeared to be the earliest progenitors of wood formation in cottonwood. It was concluded that the concept of residual meristem and its corollary, the hypothesis that acropetally developing procambial strands determine the inception sties of new primordia, apply to the cottonwood apex.

Die Entwicklungsgeschichte der Blattnervatur beim FarmStenochlaena tenuifolia (Blechnaceae)

Morphogenesis determines histogenesis both in primary leaves and in leaves of mature plants ofStenochlaena at different stages of development. The procambial strands are formed successively from the inner, more advanced parts of the leaf towards the margin, influenced by the main direction of growth. Changes in the direction of growth, e.g. from transverse to longitudinal, cause the formation of a simple network of nerves. A regular pattern originates from the differentiation of anticlinal nerves.

Floral development of Ruppia maritima var. maritima

A hyaline, unvascularized sheath envelops a portion of the inflorescence near maturity. Though resembling an appendage of the main axis, in early ontogeny it develops as a prophyll of the renewal growth apex below the inflorescence. Two flowers develop on the inflorescence axis, subopposite each other. Fertile appendages are initiated in an acropetal sequence on each floral bud. The first to form, in the median position, are the two stamens, the lower preceding the upper. Each stamen develops two bisporangiate thecae separated by a broad connective. A dorsiventral outgrowth is initiated slightly abaxially near the tip of the connective at the stage of theca differentiation. This outgrowth appears to be homologous with a similar outgrowth in Potamogeton densus, but not with the sterile appendages of the Potamogeton flower which, by some authors, have incorrectly been interpreted as connective outgrowths. Each carpel arises as a radial primordium which becomes peltate after its inception. One ovule is initiated at the adaxial portion (Querzone). The stigma becomes broad and flat, lobing at its margins. A slight outgrowth develops at the abaxial wall of the carpel. The floral apex has a two-layered tunica. The primordia of the stamens, carpels, and ovules arise by periclinal divisions in the second layer. Procambial development is acropetal following closely primordial inception. Each appendage, including the ovule, receives one procambial strand. The outgrowths of the connective and the carpel lack procambium.

Floral development of Butomus umbellatus

The primordia of the floral appendages appear in acropetal succession and develop in the order in which they appear. The primordia of each whorl of appendages are formed in a rapid sequence. After the inception of outer tepal primordia, the floral apex becomes triangular. On each angle, one inner tepal primordium together with the primordia of a pair of outer stamens and an inner stamen is formed. The triangularity of the floral apex might be interpreted as an indication of the formation of petal–stamen (CA) primordia as reported for Alisma and Hydrocleis. If this is the case, the primary pattern of organogenesis of the Butomus flower is trimerous and tetracyclic, i.e. one whorl of outer tepals, one complex of inner tepals and stamens, and two whorls of pistils. The floral apices have a two-layered tunica surrounding a central corpus. The initiating divisions in the formation of all floral appendages occur in the second tunica layer. In the case of stamen primordia, the outer corpus is also involved. Procambial development is acropetal. One procambial strand differentiates into each floral appendage shortly after its inception. Additional procambial strands are formed in the pedicel and the perianth and gynoecium. The relationships of Butomus to the Magnoliidae are discussed.

Floral development of Potamogeton densus

The floral appendages of Potamogeton densus are initiated in an acropetal sequence. The first primordia to be seen externally are those of the lateral tepals, though sectioning young floral buds (longitudinally, parallel to the inflorescence axis) reveals initial activity in the region of the lower median (abaxial) tepal and stamen at a time when the floral meristem is not yet clearly demarcated. The lateral (transversal) stamens are initiated simultaneously and unlike the median stamens each arises as two separate primordia. The upper median (adaxial) tepal and stamen develop late in relation to the other floral appendages, and in some specimens are completely absent. Rates of growth of the primordia vary greatly. Though the lower median tepal and stamen are initiated first, they grow slowly up to gynoecial inception, while the upper median tepal appears late in the developmental sequence but grows rapidly, soon overtaking the other tepal primordia. The four gynoecial primordia arise almost simultaneously, although variation in their sequence of inception occurs. The two-layered tunica of the floral apices gives rise to all floral appendages through periclinal divisions in the second layer. The third layer (corpus) is involved as well in the initiation of the stamen primordia. Procambial strands develop acropetally, lagging behind primordial initiation. The lateral stamens though initiating as two primordia each form a single, central procambial strand, which differentiates after growth between the two primordia of the thecae has occurred. A great amount of deviation from the normal tetramerous flower is found, including completely trimerous flowers, trimerous gynoecia with tetramerous perianth and androecium, and organs differentiating partially as tepals and partially as stamens.

Histology and ultrastructure of oat blue dwarf virus infected oats

The oat blue dwarf virus is a small spherical virus (28–30 nm) in diameter and is obligatorily transmitted by the aster leafhopper Macrosteles fascifrons Stål. The virus causes abnormalities in the phloem development of infected plants. Hyperplasia and limited hypertrophy of phloic procambium, in a given procambial strand, begin only after the maturation of the first protophloem sieve-tube element in that particular localized area. The majority of phloem elements produced in hyperplastic areas are parenchymatous, have truncated end walls, and lack sieve plates. Electron-microscopic observations substantiated the hypothesis that the virus was phloem-limited by revealing virus particles only in phloem elements. The greatest accumulation of virus particles was observed in the region between immature and fully vacuolated phloem elements, implicating virus synthesis in immature elements. Crystals of virus particles were often large enough to be seen by light microscopy.

Life-history ofCycas circinalis L

The paper deals with the germination of the seeds ofCycas circinalis L, the growth and structure of the different parts of the seedling in the stem, root, cotyledons and plumule. The anatomy of four different types of roots, is given. The anatomy of the stem, cotyledonary plate, procambial strands and their orientation in the stem and root are described. The origin and development of stomata found on the hypocotyledonary region of the embryo are given in detail. The cotyledons, their vasculature and final fate are described.

Ontogeny of the vegetative and floral apex of Calycanthus occidentalis

The early ontogeny of the flower of Calycanthus occidentalis is described. The pattern of initiation, procambial development, apical growth, and marginal growth of floral bracts and tepals is similar to that of the leaf, while stamens, staminodia, and carpels are not leaf-like in some aspects of their development. The initiating periclinal divisions of staminodia and carpels occur in the third and fourth layers from the surface and procambium development in the carpel is discontinuous and bidirectional. These similarities in development to some axillary buds are probably due to the position of the young primordia which are isolated from developing procambial strands by parenchyma tissue. The concave floral cup is formed through the activity of an intercalary meristem concurrently with the early stages of appendage ontogeny.

Floral development of Alisma triviale

The primordia of the floral appendages are initiated in acropetal order. They develop in the same order in which they appear but for the petals, which are retarded in their early growth and mature rapidly shortly before anthesis. While the sepal primordia are dorsiventral from their inception, the primordia of other appendages are of nearly radial symmetry and become more or less dorsiventral in their later stages of development. Each petal primordium together with the primordia of a stamen pair arise on one common petal–stamen (CA) primordium. The many pistil primordia arise on three antesepalous gynoecial bulges and the area between them. Thus, in its development the flower exhibits primarily a tricyclic trimerous plan. The floral apices have a two-layered tunica up to the stage of pistil inception. The initiation of all floral appendages occurs by periclinal divisions in the second layer. The third layer (corpus) may contribute, especially in the case of the petal–stamen primordia and the gynoecial bulges. The development of procambium is acropetal. Each primordium receives a single procambial strand shortly after its initiation. Thus, procambial differentiation occurs as a response to primordial inception and not according to the principle of the conservatism of vascular tissue. Additional procambial strands may differentiate as a response to increase in size. The relationships of Alisma to some ranalian families are discussed. Since the floral pattern of Alisma may be considered as a secondary derivation from a trimerous pattern, it does not appear primitive at all. Other primitive features such as apocarpy and lack of fusion of pistil margins are however retained. Thus, Alisma is a good example for heterobathmy.

Vascular System of the Stem of the Wheat Plant .II. Development

The sequence of vascular differentiation in the shoot of the main tiller of Triticum aestivum L. was reconstructed from seriai transverse sections of shoot apices made at various stages of development. The pattern of initiation and development of the pro- cambial strands was confirmed. The provascular bundles of the pith plexus arose independently and developed acropetally from the base of the future node. Early dif- ferentiation of proto-phloem and -xylem in the main procambial strands proceeded bidirectionally up the leaf primordia and down the stem from a site of initiation isolated from other differentiated vascular elements. Further development was basipetal from the tip of the primordia, and the rate of differentiation of the sieve elements was sufficient to maintain phloem continuity across the intercalary meristems of the laminae, sheaths, and internodes. Within the developing nodes sieve tube differentiation in the cross-linking strands lagged behind that of the leaf traces they interconnected, and this may influence the movement of photoassimilate from a recently expanded leaf to the apex.