Morphological and anatomical studies inTribulus terrestris

Seedling morphology and vascular course inTribulus terrestris were studied. This species has no erect stem, but four buds appear immediately above the cotyledonary node and grow into prostrate shoots. They were determined to be the main axis of the seedling and the axillary branches of the earliest three foliage leaves, which arise very close to each other. All the leaves, including cotyledons, are vascularized with four bundles among which two are related to a single median gap. When two leaves are attached to one node, lateral traces to the opposed leaves are derived by bifurcation of a single bundle at either side of the stem. In the shoot with a series of alternate leaves, the median pair of traces to every other leaf are found on the same orthostichy. In the branch of which the first node bears no flower but an anisophyllous pair of leaves, the smaller leaf at the node was proven to be the first prophyll because its median traces are superposed by those to the leaf at the next node.     

DEVELOPMENT AND PHYLLOTAXIS OF THE VEGETATIVE AXILLARY BUD OF MICHELIA FUSCATA

Tucker, Shirley C. (Northwestern U., Evanston, III.) Development and phyllotaxis of the vegetative axillary bud of Michelia fuscata . Amer. Jour. Bot. 50(7): 661–668. Illus. 1963.—The vegetative axillary buds of Michelia fuscala are dorsiventrally symmetrical with 2 ranks of alternately produced leaves. The direction of the ontogenetic spiral in each of these buds is related both to the symmetry of the supporting branch and to the position of the bud along the branch. On a radially symmetrical branch, all the axillary buds are alike—all clockwise, for example. But in a dorsiventrally organized branch the symmetry alternates from clockwise in 1 axillary bud to counterclockwise in the next bud along the axis. Leaf initiation and ontogeny of the axillary apical meristem conform with those of the terminal vegetative bud. The axillary bud arises as a shell zone in the second leaf axil from the terminal meristem. During this process the axillary apex develops a zonate appearance. The acropetally developing procambial supply of the axillary bud consists wholly of leaf traces. At the nodal level the bud traces diverge from the same gap as the median bundle trace of the subtending leaf. Only the basal 1–2 axillary buds which form immediately after the flowers elongate each year, while the majority remains dormant with 3 leaves or fewer.     

STRUCTURE AND DEVELOPMENT OF THE DIMORPHIC BRANCH SYSTEM OF THEOBROMA CACAO

The vegetative morphology of Theobroma cacao, the cacao tree, was studied in order to provide a foundation for further investigations on the morphogenesis of the cacao dimorphic shoot system. The seedling of cacao has a determinate orthotropic shoot with a (2+3) phyllotaxis. Branch dimorphism is initiated after 1 to 2 years of growth at which time the apical meristem of the orthotropic shoot aborts and a pseudowhorl of plagiotropic branches is initiated from axillary positions in the shoot tip. The plagiotropic branches are characterized by a distichous phyllotaxis and indeterminate growth. Subsequently an axillary bud below the pseudowhorl develops into a new orthotropic shoot. The apical meristem of this shoot eventually aborts and another pseudowhorl is formed. The apical anatomy of the two types of shoots is similar. The developmental potentiality of the orthotropic shoot axillary buds to form one or the other type of shoot was investigated. The phyllotaxis of the axillary buds of the orthotropic shoot is spiral and that of the axillary buds of the plagiotropic branch is distichous. Pruning and apical puncture experiments showed that the axillary buds of a plagiotropic branch, and of an orthotropic seedling shoot which has not yet formed a pseudowhorl, always give rise to the parent type of shoot. However, the axillary buds of an orthotropic shoot which already bears a pseudowhorl give rise to either type of shoot for several nodes below the point of origin of the pseudowhorl. The type of shoot has no influence on the form of branch which develops from an axillary bud grafted to it. This evidence supports the hypothesis that the axillary buds are initiated as one or the other type of shoot, i.e., once initiated they are predestined.     

AXILLARY AND DICHOTOMOUS BRANCHING IN THE PALM CHAMAEDOREA

In both Chamaedorea seifrizii Burret and C. cataractarum Martius each adult foliage leaf subtends one axillary bud. The proximal buds in C. seifrizii are always vegetative, producing branches (= new shoots or suckers); and the distal buds on a shoot are always reproductive, producing inflorescences. The prophyll and first few scale leaves of a vegetative branch lack buds. Transitional leaves subtend vegetative buds and adult leaves subtend reproductive buds. Both types of buds are first initiated in the axil of the second or third leaf primordia from the apex, P2 or P3. Later development of both types of bud tends to be more on the adaxial surface of the subtending leaf base than on the shoot axis. Axillary buds of C. cataractarum are similarly initiated in the axil of P2 or P3 and also have an insertion that is more foliar than cauline. However, all buds develop as inflorescences. Vegetative branches arise irregularly by a division of the apex within an enclosing leaf (= P1). A typical inflorescence bud is initiated in the axil of the enclosing leaf when it is in the position of P2 and when each new branch has initiated its own P1. No scale leaves are produced by either branch and the morphological relationship among branches and the enclosing leaf varies. Often the branches are unequal and the enclosing leaf is fasciated. The vegetative branching in C. cataractarum is considered to be developmentally a true dichotomy and is compared with other examples of dichotomous (= terminal) branching in the Angiospermae.     

LIGNOTUBER ONTOGENY IN THE CORK-OAK (QUERCUS SUBER; FAGACEAE) I. LATE EMBRYO

Changes at the cotyledonary node of the cork-oak (Quercus suber L.) were examined during the embryo maturation phase using light microscopy and scanning electron microscopy techniques. During the maturation phase the embryo axis elongates by diffuse growth, the apical meristem forms the first leaf primordia, and the radicle meristem remains inactive. The primary axis of the embryo bears, axillary to the cotyledons, in the range of five to seven pairs of lateral buds at differing stages of development. Two or three pairs of these buds are visible, occurring on the upper unfused portion of the embryonic axis, while the remaining buds are hidden by the fused cotyledonary tissues. Lateral buds develop from clusters of cells in the peripheral meristem forming a shell zone delimiting the bud meristem. Lateral buds do not undergo much development until germination begins. The results are discussed with reference to the possible role of the cotyledonary node as the source of the lignotuber in the cork-oak.     

Function of transfer cells in the nodal regions of stems,particularly in relation to the nutrition of young seedlings

Summary The distribution and time course of development of transfer cells in the hypocotyl region of lettuce (Lactuca sativa L.) and groundsel (Senecio vulgaris L.) are examined by light microscopy of serial sections through a sequence of ages of hypocotyls. Investments of xylem transfer cells occur in departing traces to the cotyledons and, later, in the traces to foliage leaves; phloem transfer cells are widely distributed but particularly prominent in those bands of protophloem in the plumule vasculature which lie alongside xylem of the cotyledonary traces. Both classes of transfer cell are well endowed with wall ingrowths before differentiation of xylem and perforation of stomata occurs in the plumule. Autoradiographic evidence is obtained of a transport pathway from cotyledonary trace xylem elements to xylem transfer cell to plumule, and analyses of xylem sap collected from above or below the zones of transfer cells in the hypocotyl show that certain materials can be removed from the xylem sap by transfer cells as it moves towards the cotyledons. From these findings it is concluded that the seedling transfer cells play an important role in nutrition of the young plumule, particularly before the latter has become adequately connected with the vascular systems of cotyledons and root.Experiments on the experimental modification of transfer cell development in the hypocotyl suggest that both photosynthetic fixation of carbon dioxide and a transpirational loss of water by a cotyledon must take place before the presumptive xylem transfer cells in its traces can develop normal sets of wall ingrowths.Discussion is extended to the general role of transfer cells in the nodal regions of stems. Possible functions envisaged are, the general nutrition of young tissues of the apical region, the abstraction of assimilates for local storage, the transfer of assimilates to axillary buds released from apical dominance, and the interchange of assimilates between adjacent vascular traces running through the node.     

DEVELOPMENTAL MORPHOLOGY OF THE EMBRYO AND SEEDLING OF RHIZOPHORA MANGLE L. (RHIZOPHORACEAE)

The embryo of Rhizophora mangle L. is initially attached to the integument by a long multiseriate suspensor. Its basal cells lyse, and intrusive growth of the endosperm envelops the embryo, forces the micropyle open, and often carries the embryo out of the integument. Thus, “germination” is effected by growth of the endosperm rather than of the embryo. The surface of the endosperm differentiates into a layer of peculiar transfer cells. The cotyledonary body initiates as a toroidal primordium, which later becomes lobed; most of the free portions ultimately fuse. After “germination,” the axis of the viviparous seedling grows by a diffuse intercalary meristem below the cotyledonary node. Before seedling abscission, the shoot apex produces three pairs of leaves, the first of which aborts, leaving the rest of the plumule protected by their stipules. The (immersed) radicle apex is nearly inactive, but lateral roots arise early in seedling development; these are usually the first or only roots to grow during establishment. Ten provascular strands “differentiate” in the cotyledons; a hollow provascular cylinder develops in the hypocotyl. Initial vascular differentiation in the latter is of many alternate poles of xylem and phloem; later, de novo differentiation of metaxylem opposite the protophloem poles, and vice versa, produces collateral bundles. Xylem maturation is endarch over most of the length of the hypocotyl, but tangential and random series of metaxylem vessels occur in the radicle end.     

ANATOMICALLY PRESERVED GLOSSOPTERIS STEMS WITH ATTACHED LEAVES FROM THE CENTRAL TRANSANTARCTIC MOUNTAINS,ANTARCTICA

Stems and buds of Glossopteris skaarensis Pigg and buds of G. schopfii Pigg from the Permian Skaar Ridge locality in the central Transantarctic Mountains, Antarctica demonstrate the first anatomically preserved glossopterids known with stem/leaf attachment. Stems of G. skaarensis are 1–12 mm in diameter ( = 3.1 mm) with a broad pith, poorly defined primary xylem, and a zone of secondary xylem up to 6 mm thick. Pycnoxylic wood conforming to Araucarioxylon Kraus is composed of tracheids with uni- to biseriate oval to hexagonal bordered pits on radial walls, uniseriate rays one to a few cells high, and cupressoid to taxodioid cross-field pitting. Stems have a narrow zone of secondary phloem, aerenchymatous cortex with scattered sclereids, and sometimes a narrow periderm. Two wedge-shaped leaf traces each bifurcate to form four strands in the base of each petiole. Small axillary branches are vascularized by double branch traces that fuse at the margin of the main axis. Buds of G. skaarensis have leaves with narrow lateral laminae and a thickened midrib containing a wide lacuna, delicate vascular strands, and a prominent hypodermis. In contrast, buds of G. schopfii have uniformly thick leaves with prominent, circular vascular bundle sheaths. These anatomical details are used to reconstruct individual types of glossopterid plants, providing new information toward understanding the ecology and evolution of this important group of Permian seed plants.     

The origin, initiation and development of axillary shoot meristems in Lotus japonicus

BACKGROUND AND AIMS: Lotus japonicus 'Gifu' develops multiple axillary shoots in the cotyledonary node region throughout the growth of the plant. The origin, initiation and development of these axillary meristems were investigated. METHODS: Morphological, histological and mRNA in situ analyses were done to characterize the ontogeny of cotyledonary axillary shoot meristems in Lotus. Morphological characterization of a putative Lotus shoot branching mutant (super-accessory branches) sac, is presented. KEY RESULTS: By using expression of an L. japonicus STM-like gene as a marker for meristematic tissues, it was demonstrated that groups of cells maintained in the meristematic state at the cotyledonary axil region coincide with the sites where additional axillary meristems (accessory meristems) form. A Lotus shoot branching mutant, sac, is a putative Lotus branching mutant characterized by increased proliferation of accessory shoots in all leaf axils including the cotyledons. CONCLUSION: In Lotus, axillary shoot meristems continually develop at the cotyledonary node region throughout the growth of the plant. These cotyledonary primary and accessory axillaries arise from the position of a meristematic zone of tissue at the cotyledonary node axil region.     

SYLLEPTIC BRANCHING IN MYRSINE FLORIDANA (MYRSINACEAE)

Myrsine floridana produces all of its vegetative branches, other than those resulting from pruning or damage, by syllepsis, i.e. by the continuous development of an axillary meristem into a branch without an intervening stage of rest. These sylleptic branches, produced in series, have long and conspicuous hypopodia, broad pith connections with the parent axis, and expanded prophylls. Bud dormancy may be imposed when an axillary meristem is in the axil of the sixth or seventh youngest leaf of the parent shoot. Such axillary meristems may remain at the bud stage with only two pairs of scalelike leaves but these may later give rise to inflorescences or proleptic branches. Proleptic branches lack hypopodia, have narrow piths at their bases, and a series of leaves transitional from the original prophylls to normal foliage leaves within about ten leaves. Myrsine floridana has cortical bundles in the stem, related to the formation of minor lateral leaf traces. The hypopodia of sylleptic branches, since they are leafless, do not have cortical bundles.     

TENDRILS OF PASSIFLORA FOETIDA: HISTOGENESIS AND MORPHOLOGY

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

Branch development in Lupinus angustifolius L.#II. Relationship with endogenous ABA, IAA and cytokinins in axillary and main stem buds

The concentrations of indole-3-acetic acid (IAA), cytokinins (CK) and abscisic acid (ABA) were measured in buds of different regions (main stem and lateral branches) of Lupinus angustifolius L. (cv. Merrit) and at different stages in the development of branches. In lupin, branching patterns are the result of discrete regions of axillary branches (upper, middle and basal) which elongate at much different rates. Early in development only the main shoot elongates, followed usually by basal branch growth and then rapid upper branch growth. Branches in the middle of the main stem grow only weakly or fail to develop. Levels of IAA were generally high in the apical buds of slowly growing branches and low in buds from strongly growing branches, whereas CK levels showed the opposite relationship. CK:IAA ratio showed a closer relationship with the rate of growth of a particular branch better than the levels of either CK or IAA alone. During early stages of growth ABA concentration did not follow the rate of branch growth. However, later in development, where growth did not closely match the ratio of CK:IAA, ABA level showed a strong negative relationship with growth. A significant decrease in ABA was associated with continued strong growth of the main stem apex following a decline in CK:IAA ratio. Overall, the best relationship between the level of growth factors in apical buds and branching pattern in lupin was the ratio of CK:IAA, implying that high CK:IAA at a given bud would promote growth. ABA level appeared to play a secondary role, as a growth inhibitor.     

CONTROL OF SHOOT-RHIZOME DIMORPHISM IN THE WOODY MONOCOTYLEDON,CORDYLINE (AGAVACEAE)

In Cordyline terminalis negatively geotropic leafy shoots and positively geotropic rhizomes develop from single axillary buds on either shoots or rhizomes. All axillary buds have similar morphogenetic potential when released from apical dominance. Experiments in which the orientation of the apex is changed, organs removed, or growth regulators applied indicate that after a rhizome is initiated, it is maintained as a rhizome by auxin originating in the leafy shoot. When auxin levels are lowered by changes in the orientation of the axis or shoot removal, the rhizome apex becomes a shoot apex, which appears to be the stable state of the actively growing apex. Benzyl adenine when applied exogenously to the apex or lateral buds has the same effect as lowering the auxin level. Gibberellic acid has no effect on the apex or lateral buds. High levels of exogenous naphthaleneacetic acid cause bud release and development of rhizomes from previously inhibited axillary buds of the shoot. However, it was not possible to convert a shoot apex into a rhizome apex by auxin treatment. It is suggested that the release of buds on the lower side of horizontal branches and of buds directly above a stem girdle is caused by high auxin levels on the lower side or distal to the girdle. The experimental results are discussed in relation to naturally occurring shoot-rhizome dimorphism.     

STUDIES ON THE ONTOGENY OF THE DWARF MISTLETOES,ARCEUTHOBIUM. III. DEVELOPMENT OF THE INFLORESCENCE

In vascular plants, the apical meristem of the shoot normally represents a continuation of growth in the apical meristem of the embryo itself. This is not the case in Arceuthobium. Here the shoot apex of the embryo is rudimentary and eventually dies after infection of the host occurs. The inflorescence of Arceuthobium is, therefore, an adventitious structure originating in the endophytic system rather than from the shoot apex of the seedling. Inflorescence buds arise in either of 2 ways. In some species (A. douglasii and A. americanum), buds first appear as small meristematic protuberances on the outer surface of cortical strands. In other species (A. campylopodum), the buds arise at the ends of short branches. The former, or diffuse, type gives rise to inflorescences along the entire surface of the host branch; in the latter, or condensed, type inflorescences are formed in clusters. Early ontogeny of the inflorescence apex of both types is described. Studies of subsequent growth of the inflorescence apex show 5 well-defined plastochronic stages: (1) maximal area stage; (2) minimal area stage; (3) early post-minimal stage; (4) late post-minimal stage; and (5) pre-maximal stage.     

The development of Archaeopteris: new evolutionary characters from the structural analysis of an Early Famennian trunk from southeast Morocco

A 5 m long trunk of a young Archaeopteris/Callixylon erianum tree from the Late Devonian of Morocco shows new branching patterns for early lignophytes. This progymnosperm tree produces a helical pattern of traces that we infer belonged to reduced, short-lived, primary (apical) branches (type A) as well as two types of adventitious traces (types B and H). We infer that type-B traces supplied branches that initiate close to the site of attachment on the trunk of some, but not all type-A branches in an irregular but nonrandom pattern. Unlike ephemeral type-A branches, those of type B persist and become long-lived, potentially permanent units of the architecture of Archaeopteris trees. Type-H adventitious traces are also short-lived and occur singly or in serial groups, but differ from traces of either type A or B branches by lacking differentiation into a readily identifiable organ category. We interpret type-H traces as supplying latent primordia that could develop into either adventitious roots or shoots depending on extrinsic factors. Our new data suggest that Archaeopteris had a wide range of branch primordium amplitude. Type-B branches compare with axillary lateral branch buds of some Early Carboniferous spermatophytes (Calamopitys) and are a major developmental departure from the strictly apical, pseudomonopodial shoot branching of older aneurophyte progymnosperms. Type-H traces suggest that Archaeopteris trees had some potential for formation of adventitious roots or shoots in response to environmental factors, such as partial burial by overbank sedimentation. Collectively, these novel methods of tree branching may partly explain the extraordinary success and worldwide dominance of Archaeopteris forests on fluvially dominated, Late Devonian floodplains.     

Relationships among shoot sinks for resources exported from nodal roots regulate branch development of distal non-rooted portions of Trifolium repens L

Two manipulative experiments tested hypotheses pertaining to the correlative control exerted by nodal roots on branch development of the distal non-rooted portion of Trifolium repens growing clonally under near-optimal conditions. The two experiments, differing in their pattern of excision to manipulate the number of branches formed at the first 9-10 phytomers distal to the youngest nodal root, each found that after 20 phytomers of growth the total number of lateral branches formed on the primary stolon remained between five and seven regardless of where the branches formed along the stolon. Additional treatments established that nodal roots influenced branch development via relationships among shoot sinks for the root-supplied resources rather than through variation in the supply of such resources induced by fluctuations in photosynthate supply to roots from branches. Regression analysis of data pooled from treatments of both experiments confirmed that shoot-sink relationships for root- supplied resources controlled the branching processes on the non-rooted portion of plants. A disbudding treatment, which removed all the apical and axillary buds present on basal branches, but left other branch tissues intact, increased branch development of the apical region in the same way as did complete excision of the basal lateral branches. The apical buds and the elongation processes occurring immediately proximal to the buds were thus identified as strong sinks for the root-supplied resources. Such results suggest that branch development on the non-rooted shoot portion distal to the youngest nodal root is regulated by competition among sinks for root-derived resources, of limited availability, necessary for the processes of elongation of axillary buds and the primary stolon apical bud.     

The Effects of Temperature and Other Variables on the Contribution of Lateral Branches to Yield in Phaseolus vulgaris L.

At low temperatures (15/15 °C day/night) in controlled environmentsthe growth of lateral branches at the cotyledonary node ofPhaseolusvulgarts L is suppressed The suppression can be overcome byraising the temperature of the buds by approximately 6 °Cwith small electric heaters In order to test the practical significanceof the induction of branching for yields of the commercial crop,seedlings were raised in contrasted regimes and then transplantedto the field The effects of pretreatment regime on final yieldwere small, changes in the yield from axillary branches tendedto be balanced by compensatory changes in the yield from themain stem In another field experiment, synthetic growth substanceswere applied in order to suppress or enhance branching Changesin the amount of yield carned on branches were again offsetby compensatory changes in the yield from the main stem Compensatoryeffects between branches and main stem were also found in avariety trial However, in an experiment on a single cultivarand various levels of N fertilizer, compensatory effects werenot found, here, branch and main stem yields were positively,rather than negatively, correlated These results are discussedin relation to the intrinsic factors that govern yield in Pvulgaris Phaseolus vulgaris L, dwarf bean, axillary branches, correlative inhibition, temperature, growth substances, plant density, yield     

Influence des racines sur le développement végétatif ou floral des bourgeons cotylédonaires chez le Scrofularia arguta: role possible des cytokinines

Influence of roots on the vegetative or floral development of cotyledonary buds of Scrofularia arguta Sol.: A possible cytokinin role. This study shows that the presence of “nonabsorbing roots” insures a vegetative development of cotyledonary buds cultured in vitro whereas buds growing without roots produce flowers early. In the same way, roots suppress floral expression of axillary meristems of the same cotyledonary buds and induce these buds to vegetative functioning. Various trophic modifications in the culture medium are ineffective on non-rooted buds as also are gibberellin As and adenine. On the contrary, several cytokinins (kinetin, benzyladenine and zeatin) exert the same influence as roots. These results suggest that roots regulate meristematic functioning through cytokinins.     

THE ORIGIN AND DEVELOPMENT OF INDUCED BRANCHES IN HELIANTHUS

The development of buds and their vascular connections are described for Helianthus annuus and H. bolanderi. Bud meristems of H. annuus usually become isolated by parenchymatization of the bud traces in the cortical zone. If the buds are induced to grow as a result of decapitation of the terminal meristem, a continuous range between typical primary connections and pseudo-adventitious connections are made between the main axis and the lateral buds. Considerable growth of the branches occurs within 48 hours after decapitation. Axillary buds of H. bolanderi grow continuously, and both the meristem and vascular system of the buds are derived directly from the apical meristem of the shoots.     

An in vitro technique for the production de novo of multiple shoots in cotyledon explants of cucumber (Cucumis sativus L.)

A technique is described for the production de novo of cucumber (Cucumis sativus L.) shoots in the presence of cytokinin using cotyledon explants. The shoots, which arose from adventitious buds and not from enhanced axillary branching, are confined to a specific region at the base of the cotyledon. Concentrations (4 mgl^–1 or less) of the cytokinins 6-benzylaminopurine, kinetin and N⁶-(2-isopentenyl)adenine, are all effective in producing adventitious buds. It is possible to achieve a yield of 23 shoots per cotyledon by removal of the axillary bud. The yield is increased to 50 shoots per cotyledon by cutting the basal region of the cotyledon into small pieces prior to culturing. These techniques may be useful for transformation studies in cucumber.