DIFFERENTIATION OF LATERAL SHOOTS AS THORNS IN ULEX EUROPAEUS

Ulex europaeus is a much-branched shrub with small, narrow, spine-tipped leaves and axillary thorn shoots. The origin and development of axillary shoots was studied as a basis for understanding the changes that occur in the axillary shoot apex as it differentiates into a thorn. Axillary bud primordia are derived from detached portions of the apical meristem of the primary shoot. Bud primordia in the axils of juvenile leaves on seedlings develop as leafy shoots while those in the axils of adult leaves become thorns. A variable degree of vegetative development prior to thorn differentiation is exhibited among these secondary thorn shoots even on the same axis. Commonly the meristems of secondary axillary shoots initiate 3–9 bracteal leaves with tertiary axillary buds before differentiating as thorns. In other cases the meristems develop a greater number of leaves and tertiary buds as thorn differentiation is delayed. The initial stages in the differentiation of secondary shoot meristems as thorns are detected between plastochrons 10–20, depending on vigor of the parent shoot. A study of successive lateral buds on a shoot shows an abrupt conversion from vegetative development to thorn differentiation. The conversion involves the termination of meristematic activity of the apex and cessation of leaf initiation. Within the apex a vertical elongation of cells of the rib meristem initials and their immediate derivatives commences the attenuation of the apex which results in the pointed thorn. All cells of the apex elongate parallel to the axis and proceed to sclerify basipetally. Back of the apex some cortical cells in which cell division has persisted longer differentiate as chlorenchyma. Although no new leaves are initiated during the extension of the apex, provascular strands are present in the thorn tip. Fibrovascular bundles and bundles of cortical fibers not associated with vascular tissue differentiate in the thorn tip and are correlated in position with successive incipient leaves in the expected phyllotactic sequence, the more developed bundles being related to the first incipient leaves. Some secondary shoots displayed variable atypical patterns of meristem differentiation such as abrupt conversion of the apex resulting in sclerification with limited cell elongation and small, inhibited leaves. These observations raise questions concerning the nature of thorn induction and the commitment of meristems to thorns.     

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.     

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.     

The occurrence of shoot teratomata in tissue cultures of Taraxacum officinale

Summary Tissue cultures of Taraxacum officinale have been isolated showing either normal organogenesis, callus growth only, or teratological shoots. The latter are apparently stable and the teratomata range from tumerous outgrowths, to flattened thalloid forms and short shoots with strap shaped leaves. The proximal regions of such shoots are tumerous and no distinction between apex, leaf and bud primordia is possible.     

Influence of benzyladenine and gibberellic acid on organogenesis in lsquo;Crimson Giantrsquo; Easter cactus

Experiments were performed to determine the influence of gibberellic acid (GA₃) and benzyladenine (BA) on organogenesis of lsquo;Crimson Giantrsquo; Easter cactus [Hatiora gaertneri (Regel) Barthlott] phylloclades cultured in vitro. The numbers of flower buds and new phylloclades increased linearly as BA concentration increased from 0 to 444.1 micro;M. GA₃ increased the number of new phylloclades when present in moderate concentrations (2.9 or 28.9 micro;M), but inhibited flower bud formation when present in concentrations as low as 0.3 micro;M. The inhibitory effect of GA₃ on flower bud formation was diminished when the medium was amended with BA at 44.4 or 444.1 micro;M. Explants cultured in media that contained 288.7 micro;M GA₃ produced fewer organs (new phylloclades plus flower buds) compared to those cultured in media with 0, 0.3, 2.9, or 28.9 micro;M GA₃. BA and GA₃ concentrations also affected the percentage of explants with flower buds and the percentage of explants with new phylloclades. This study shows that organogenesis in H. gaertneri can be controlled by varying the concentrations of BA and GA₃ in the culture medium.     

The influence of expanding leaves and the reproductive stem apex on apical dominance in Lolium multiflorum

In vegetative plants of Lolium multiflorum removal of the two youngest emerging leaves resulted in increased expansion of basal tiller buds. A similar release of inhibition of tiller buds took place if the floriferous apex was removed. The surgical procedures did not affect the response. Under conditions of N-deficiency total tiller number was reduced but on removal of the apex the deficient plants showed an increased initial rate of tiller bud expansion. Apical dominance during the vegetative stage of growth in this grass was apparently due to the expanding leaves in the vegetative apex, but in the flowering plant the control was exerted by the inflorescence or the elongating stem.     

SCLEREID DISTRIBUTION IN THE LEAVES OF PSEUDOTSUGA UNDER NATURAL AND EXPERIMENTAL CONDITIONS

Al -talib , Khalil H., and John G. Torrey . (U. California, Berkeley.) Sclereid distribution in the leaves of Pseudotsuga under natural and experimental conditions. Amer. Jour. Bot. 48(1): 71–79. Illus. 1961.—A study of the distribution of sclereids in cleared leaves taken from 1-, 2-, and 4-year-old shoots of an adult tree of Pseudotsuga menziesii (Mirb.) Franco showed a repeated pattern of sclereid distribution along the shoot axis with many sclereids in the basal leaves grading into few or no sclereids in the terminal leaves of each year's growth. Attempts were made to influence sclereid distribution by bud defoliation of attached branches with and without auxin treatment and by testing the effects of growth-regulating substances on sclereid formation in leaves of excised buds of Pseudotsuga cultured in vitro. Whereas removal of the basal ¾ of the leaves at the time of bud unfolding had no effect on bud, leaf or sclereid development, removal of the leaves of the upper half or complete defoliation led to premature expansion of next year's terminal bud with leaves developing in part from presumptive bud-scale primordia. Indoleacetic acid at 0.5% in lanolin paste applied to the defoliated region prevented this premature bud expansion. Defoliation of the basal half did not affect sclereid formation in the terminal leaves. Sclereid development in leaves of prematurely expanded buds on defoliated branches was normal except in the few cases where bud expansion occurred in the presence of low-auxin concentrations. Then, sclereid development was inhibited. Sclereid formation in leaves of excised buds grown in nutrient culture was generally much less frequent than in intact branches, and auxin treatment still further reduced the frequency of sclereids. It was concluded that sclereid initiation and differentiation in the intact plant may well be under the control of hormonal factors in the plant, one of which may be auxin.     

LEAF-OPPOSED BUDS IN MUSA: THEIR DEVELOPMENT AND A COMPARISON WITH ALLIED MONOCOTYLEDONS

A single, lateral, vegetative bud which is positioned 180° from the axil of a leaf is a generic feature of Musa (Musaceae). Such leaf-opposed buds occur in all ten species and five cultivars examined, representing all four sections of the genus and all groups of cultivated bananas and plantains. The bud arises relatively late and is first visible as a vascular-free “clear zone” in the axis directly below the future bud meristem site. It is first associated with the fifth or sixth leaf primordium from the apex. A defined superficial meristem develops on the stem directly above the insertion of the leaf margins one or more plastochrons later. Normal, basically axillary, vegetative buds occur in the closely related genera: Orchidantha (Lowiaceae), Heliconia (Heliconiaceae), Strelitzia, and Ravenala (Strelitziaceae). These buds arise in the axil of the first to the third leaf primordium in a manner similar to most other monocotyledons. Axillary vegetative buds also occur in the remaining families of the Zingiberales: Cannaceae, Costaceae, Marantaceae, and Zingiberaceae.     

SYMMETRY AND DEVELOPMENT OF BUTOMUS UMBELLATUS (BUTOMACEAE) AND LIMNOCHARIS FLAVA (LIMNOCHARITACEAE)

The prostrate rhizome of Butomus umbellatus produces branch primordia of two sorts, inflorescence primordia and nonprecocious vegetative lateral buds. The inflorescence primordia form precociously by the bifurcation of the apical meristem of the rhizome, whereas the non-precocious vegetative buds are formed away from the apical meristem. The rhizome normally produces a branch in the axial of each foliage leaf. However, it is unclear whether the rhizome is a monopodial or a sympodial structure. Lateral buds are produced on the inflorescence of B. umbellatus either by the bifurcation or trifurcation of apical meristems. The inflorescence consists of monochasial units as well as units of greater complexity, and certain of the flower buds lack subtending bracts. The upright vegetative axis of Limnocharis flava has sympodial growth and produces evicted branch primordia solely by meristematic bifurcation. Only certain leaves of the axis are associated with evicted branch primordia and each such primordium gives rise to an inflorescence. The flowers of L. flava are borne in a cincinnus and, although the inflorescence is simpler than that of Butomus umbellatus, the two inflorescences appear to conform to a fundamental body plan. The ultimate bud on the inflorescence of Limnocharis flava always forms a vegetative shoot, and the inflorescence may also produce supernumerary vegetative buds. Butomus umbellatus and Limnocharis flava exhibit a high degree of mirror image symmetry.     

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.     

ORIGIN AND EARLY MORPHOGENESIS OF LATERAL BUDS IN THE FERN DAVALLIA

The general organography, vascular organization, and leaf and bud development in Davallia solida and D. trichomanoides are described. These epiphytic species have creeping shoots with dorsally-borne leaves in a distichous phyllotaxis and the buds occur near each leaf base. Roots are borne on the ventral and flanking surfaces of the rhizome, but only at bud positions. The vascular pattern of these species is a perforated solenostele. Leaf and bud traces have distinctly different origins. While the proximity of buds to leaves has suggested that bud origin is axillary, observations show that the origin of buds is cauline and that their position is extra-axillary from inception. The stages of structural morphogenesis in Davallia buds differ significantly from the scheme proposed by Wardlaw. The principal difference is the absence of a resting period occurring between the origin and continued development of buds in Davallia. The elongated internodes which separate leaf-bud pairs from one another, the topographically distinct and predictable positions of leaves and buds, the structural equivalence of unexpanded buds, and vascular differences in leaves and buds make Davallia an useful species for physiological studies of differential bud expansion.     

Abscissic acid localization by light microscopic immunohistochemistry in Chenopodium polyspermum L. Effect of water stress

An indirect immunohistochemical technique was developed using a rabbit anti-abscissic acid (ABA) serum and the soluble peroxidase-antiperoxidase (PAP) complex for the localization of endogenous ABA in the aerial parts of Chenopodium. Terminal bud, axillary bud bearing nodes, and adult leaves were prefixed by a soluble carbodiimide to obtain the coupling of ABA on cellular proteins and postfixed by a conventional mixture of aldehydes. They were then embedded in paraffin or in plastic. Numerous controls were carried out on sections and on a model system to test the validity of the technique. Based on the staining patterns observed along the plant, an apico-basal gradient of ABA was revealed. In the older buds, ABA was mainly concentrated in the quiescent meristematic cells of the apex. Phloem cells of the main axis and chloroplasts of the leaves were specifically labeled. No reaction product was visualized in the parenchyma cells or in the cambial zone. Water stress, which is known to increase ABA content, induced an increase of immunoreactivity within the same compartments. This physiological test validates the stain.     

THE DEVELOPMENTAL ANATOMY OF LONG-BRANCH TERMINAL BUDS OF PINUS BANKSIANA

A study of the composition of long-branch terminal buds (LBTB) of Pinus banksiana Lamb. and the yearly periodicity associated with their formation, development, and elongation was undertaken. Each LBTB has lateral bud zones and zones of cataphylls lacking axillary buds. When present, staminate cone primordia differentiate from the lowest lateral buds in the lowest lateral bud zone of the LBTB. Ovulate cone primordia and lateral long-branch buds can differentiate from the upper lateral buds in any lateral bud zone. When both types of buds are present, lateral long-branch buds are uppermost. Dwarf-branch buds occur in all lateral bud zones. During spring LBTB internodes elongate, new cataphylls are initiated, dwarf branches elongate, needles form and elongate, pollen forms and is released, and ovulate cones are pollinated. During summer buds form in the axils of the newly formed cataphylls. By early fall the new LBTB are in overwintering condition and the four types of lateral buds are discernable. The cytohistological zonation of the LBTB shoot apex is similar to that of more than 20 other conifer species. Cells in shoot apices of pine are usually arranged in distinct zones: apical initials, subapical initials, central meristem, and peripheral meristem. Periclinal divisions occur in the surface cells of the apex; therefore no tunica is present. At any given time, shoot apex volume and shape vary among LBTB in various positions on a tree. In any one LBTB on a tree, shoot apex shape changes from a low dome during spring to a high dome during summer to an intermediate shape through fall and winter.     

A COMPARATIVE DEVELOPMENTAL STUDY OF THE MUTANT SIDESHOOTLESS AND NORMAL TOMATO PLANTS

'Sideshootless,’ a mutant strain of tomato which does not produce axillary buds during vegetative growth, was compared with normally branching plants in order to study the nature of development particularly with regard to axillary buds. Sectioned material revealed no indication of axillary bud initiation in the sideshootless plant at any time during the vegetative phase of growth. In the normal plants, buds were noted to arise in the axil of the fifth youngest leaf. The buds take their origin in tissue which is in direct continuity with the apical meristem. The bud primordia later become set apart from the apex as vacuolation takes place in the surrounding tissue. At the time of floral initiation, the mutant and normal strains behave similarly. Axillary buds appear in the axils of the 2 leaves immediately below the floral apex. One of the buds elongates to overtop the existing plant axis; the other develops as a typical sidebranch. The inflorescence is pushed aside in the process. This pattern is repeated with each inflorescence; thus an axis composed of several superimposed laterals results.     

The Initiation and Growth of Axillary Bud Primordia in Relation to Flowering in Trifolium repens L.

The initiation and growth of axillary bud primordia in relationto the growth of their subtending leaves was observed at theapices of three clones (A. B. and C) of white clover grown invarious combinations of photoperiod and temperature. ClonesA, B, and C flower in response to low temperatures, and clonesA and C, but not B, in response to a transfer from short tolong photoperiods at higher temperatures. The rate of growth of buds and leaves from node to node waslittle influenced by the various treatments imposed, but theinitiation of axillary bud primordia relative to the apicaldome was stimulated in conditions conducive to flowering. The number of budless leaf primordia at the apex ranged froma maximum average of 2.25 at 20° C. to approximately o.8oat 10° C. in all three clones. At the higher temperatures,runners possessed 2.06 budless nodes in short days but only1.12 in long days in clones A and C. In clone B, daylength didnot influence bud initiation at the higher temperature. The results provide evidence of the homology between vegetativeand repro-ductive axillary bud primordia. It is suggested thatflowering is brought about by the removal of an inhibition withinthe apex which leads to the precocious initiation of axillarybud primordia. Following the initiation of axillary bud primordia, the resultsshow their growth to be uninhibited for 6-7 plastochrons. Rapidinflorescence development occurs during this phase. Apical dominancehas no apparent influence on vegetative axillary buds untilthe onset of rapid petiole elongation in their subtending leaves.     

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.     

Quantitative analyses of shade-shoot architecture of conifers native to New Zealand

Shoot architecture was quantified by measuring the "maximum silhouette area ratio" (R_max). R_max was calculated from the maximum silhouette area (or projected area) of the intact shoot, divided by the silhouette area of the leaves or phylloclades (leaf-like flattened stems) when they are removed from the shoot and laid out flat. Like conifers of the Northern Hemisphere (NH) with non-appressed foliage, the R_max of shade-adapted shoots ranged from 0.5 to 1.0 in New Zealand (NZ) conifers with non-appressed foliage. Defining a "leaf" to mean either a true leaf or a phylloclade, the following was found: leaf area/leaf dry weight, leaf area/shoot dry weight, and leaf dry weight/shoot dry weight, were all similar in the shade-shoots of NZ and NH conifers. None of these variables were significantly correlated with R_max in the NZ conifers, unless species with leaves averaging less than 4 mm² in size were excluded from the analyses. Foliage dry weight/shoot projected area was strongly correlated with R_max. NZ conifers had both smaller and larger mean leaf sizes in comparison to NH conifers. The mean projected area per shade-adapted leaf of NZ conifers varied from 2.7 to 436 mm². In NH conifers, the mean projected area per shade leaf varied from 12 to 83 mm². Except for the strikingly larger range in leaf size in NZ conifers, the data support a hypothesis of strong convergent evolution of shade-shoot architecture in NZ and NH conifers. The results are discussed in relation to photosynthesis, stand production, and the ecological distribution of conifers.     

In nature dormant buds and in vitro dormant-like buds ofFraxinus excelsior L.

Summary Microcuttings ofFraxinus excelsior, sampled from adult trees during the period of cell cycle blockage of bud in G_0–1, developed long rejuvenated sprouts on the Woody Plant Medium (WPM) with benzylaminopurine (BAP) (4.0 mg/l) and indolebutyric acid (IBA) (0.03 mg/l). These sprouts had the ability to enter a resting period, building dormant-like buds when maintained on the original WPM. Sprouts developed from subcultures also entered a resting period without any transfer. Comparison of in nature buds in active growth and dormancy with buds of growing sprouts and in vitro dormant-like buds revealed similarity in behaviour at the shoot apical level. In particular, in dormant-like buds in a constant environment, shoot apical functioning was suppressed while the cell cycle of the shoot apex was blocked at the G_0–1 phase, like in nature dormant buds.Abbreviations a.u. arbitrary unit - BAP benzylaminopurine - CF cumulative frequency - d₁, d₂ diameters of the shoot apex - IBA indolebutyric acid - P_n last opposite primordia of range n - WPM woody plant medium     

INDETERMINATE GROWTH OF LEAVES IN GUAREA (MELIACEAE): A TWIG ANALOGUE

Leaves of seed plants are generally characterized as organs of determinate growth. In this regard, Guarea and related genera seem unusual in that the pinnately compound leaves of these plants contain a bud at their tip from which new pinnae expand from time to time. Previous studies (based upon superficial examinations of leaf-tip buds) have produced contradictory conclusions regarding how long the leaf apex remains meristematic and produces new pinna primordia. In order to determine whether leaf development in Guarea is truly indeterminate, we microscopically examined leaf-tip buds of G. guidonia and G. glabra. In both species, the leaf apex remains meristematic and continues to produce new pinna primordia as the leaf ages. Unexpanded leaves of G. guidonia contained an average of 23 pinna primordia, while the oldest leaves we examined had initiated an average of 44 total pinnae. In G. glabra, unexpanded leaves contained 8 pinnae, whereas an average of 28 pinnae had been initiated on the oldest leaves. These results indicate that leaf development in Guarea is truly indeterminate. Periodic examination of individual intact leaves indicated that the leaves commonly continue their growth for 2 or more years (observed maximum = 51 months). As new leaflets are initiated at the shoot apex (and subsequently expand in rhythmic flushes), older (basal) leaflets may abscise. In addition, the petiole and rachis of the leaf thicken and become woody as a result of the activity of a vascular cambium. Guarea leaves therefore seem to function as the analogue of a typical twig (stem) in general habit as well as in their indeterminate apical growth and secondary thickening.