Non-axillary branching in the palms Eugeissona and Oncosperma (Arecaceae)

FISHER, J. B., GOH, C. J. & RAO, A. N., 1989. Non-axillary branching in the palms Eugeissona and Oncosperma (Arecaceae). The south-east Asian palms, Eugeissona (Calamoideae) and Oncosperma (Arecoideae) are multiple-stemmed. The morphology and development of branching in two species of each genus were examined in Singapore, Borneo, and the Malay Peninsula. Cultivated seedling and adult plants of 0. tigillarium were also observed in Florida. A new shoot arises most often from a longitudinal abaxial groove at the base of an enclosing leaf sheath. In some instances, especially in E. tristis , the enclosing leaf has two equal, adjacent grooves such that any distinction between an original mother shoot and a lateral daughter shoot is impossible. No axillary buds occur in Eugeissona which is hapaxanthic. In Oncosperma , which is pleonanthic, axillary buds are absent from young pre-flowering stems, but an inflorescence bud occurs in the axil of each leaf in older aerial stems. Early ontogenetic stages of vegetative branching, as seen in sectioned apices, indicate that a new vegetative shoot is present on the abaxial base of the first (youngest) leaf primordium. There is no ontogenetic evidence for the displacement of an originally axillary meristem as previously described for the palm Salacca (Calamoideae). Shoot development in Eugeissona is interpreted as a putative dichotomy of the apical meristem in which the meristem centres commonly develop unequally. In Oncosperma the smaller sucker bud meristem may be described as an abaxial leaf base bud, but ontogenetic variations indicate this form of branching is close to dichotomous branching. These new examples of non-axillary branching are compared to similar cases previously reported for palms and other monocotyledons.     

Development of Axillary and Leaf-opposed Buds in Rattan Palms

Axillary vegetative buds are present in Calamus, Ceratolobus,and Plectocomiopsis. Two species of Daemonorops Sect. Piptospathaalso have axillary vegetative buds. All species of Daemonoropshave only displaced adnate axillary inflorescence buds. A singlebud is initiated in the axil of the first or second leaf primordiumin a way similar to that for axillary inflorescence buds. Themeristem is displaced during development on to the internodeabove and sometimes on to the base of the leaf above. Leaf-opposedvegetative buds occur in five species of Daemonorops Sect. Cymbospathaand in one species of Daemonorops Sect. Piptospatha. This typeof bud is initiated 180° away from the axil of the firstor second leaf primordium. It is not a displaced axillary bud,but does become adnate to the internode above like the axillarybuds. One or more leaves, transitional between juvenile andadult, on a shoot often subtend both types of buds. Myrialepishas leaf-opposed vegetative buds, but their development wasnot observed. Korthalsia has buds that are displaced about 130°from the leaf axil and are intermediate between the axillaryand the leaf-opposed condition. Other forms of vegetative budsare described: multiple buds in Plectocomia, aerial forkingin Korthalsia, and suckering from inflorescences and from aerialstems in Calamus. bud development, rattan palms, palm taxonomy, branching     

Unusual branch development in the palm, Chrysalidocarpus

Vegetative branch buds of C. lutescens are non-axillary and occur within an abaxial, tubular extension of the leaf sheath, either at the base of a shoot or aerially, a position unusual for palms. Buds are initiated on the abaxial surface of a leaf during its first plastochron (the youngest leaf primordium). The foliar origin of the vegetative bud appears to be unique for angiosperms. In contrast, inflorescence buds are axillary and are initiated as an adaxial ridge on the base of a leaf during its third plastochron (the third primordium from the apex). Aerial branches and basal suckers are developmentally identical and changes in their phyllotaxis are described. As far as can be established by comparative morphology, other species of Chrysalidocarpus have the same type of branch development as in C. lutescens. The development of branches is related to the morphogenetic characteristics of arborescent monocotyledons.     

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.     

Comparative morphology and development of inflorescence adnation in rattan palms

The morphology and development of inflorescences in 14 genera and 52 species of rattans and related genera of Lepidocaryoid palms were examined. Inflorescences are free (not adnate) in Ancistrophyllum, Eremospatha and Oncocalamus. Adnation between the inflorescence and internode above occurs in Korthalsia, Myrialepis, Plectocomia and Plectocomiopsis. Adnation between the inflorescence and both the internode and leaf sheath above occurs in Calamus, Calospatha, Ceratolobus and Daemonorops. This leaf-borne, but initially axillary, bud is displaced on to the base of the next younger leaf primordium by the second plastochrone. Later elongation of the internode further separates the inflorescence from its original node. Stages of initiation and early development of adnate buds are illustrated for ten species. Vegetative buds of some Calamus species develop like inflorescence buds. However, other species have unusual bud positions which cannot be interpreted at present. The degree of inflorescence adnation tends to be greater in presumably specialized species than in unspecialized ones.     

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.     

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.     

A morphological study of the development of the second inflorescences in strawberry (Fragaria×ananassa Duch.)

To clarify the timing of the differentiation of the first and second inflorescences in strawberry (Fragaria × ananassa Duch.), morphological changes on shoot apices during short day and low night temperature treatments were observed by scanning electron microscopy (SEM) and optical microscopy. Axillary buds just below the first inflorescence (axillary bud 1) became visible when sepal primordia of the primary flower were differentiated. By this time, other axillary buds had already developed. Axillary bud 1 developed four leaf primordia, and then a differentiated inflorescence at its summit. The phase transition of shoot apices from the vegetative to the reproductive phase may therefore trigger the differentiation of axillary bud 1 which is destined to develop into extension crowns.     

Developmental study onHypolepis punctata (Thunb.) Mett.

The third petiolar bud ofHypolepis punctata appears on the basiscopic lateral side of the petiole above the fairly developed first petiolar bud. This investigation clarified the fact that the third bud is formed neither by the activity of the meristem of the first bud nor by the meristem directly detached from the shoot apical meristem, but is initiated in the cells involved in the abaxial basal part of the elevated portion of the leaf primordium. Thus the third bud is of phyllogenous origin. This investigation further revealed that the cells to initiate the third bud are originally located in the abaxial side of the leaf apical cell complex like the cells to initiate the first bud, but are not incorporated into the meristem of the first. After the first, second and third petiolar buds have been initiated, they are carried up into fairly high regions on the petiolar base by the intercalary growth which occurs in the leaf base below the insertion level of the first and the second buds.     

Development of the epiphyllous inflorescence of Phyllonoma integerrima (Turcz.) Loes.: implications for comparative morphology*

The inflorescence primordium of Phyllonoma integerrima (Turcz.) Loes. is initiated on the adaxial side of the leaf primordium. At about the same time, a vegetative bud is formed at the base of the same leaf primordium. The vascular anatomy is the same in the fertile and sterile leaves, except that in the fertile leaf an inflorescence trace departs from the midvein of the leaf at the point where the inflorescence is inserted. Neither the inception nor the procambial supply of the inflorescence provide evidence of “congenital fusion”of inflorescence and leaf. It is also argued that the idea of an “adventitious”origin of the inflorescence is not useful in this case. Consequences for our conception of shoot construction are pointed out. It is argued that positional changes in the initiation of organs is an evolutionary process that may have remarkable effects on plant construction.     

Development of shoot inHydrangea macrophylla I. Terminal and axillary buds

The structure of shoots, in particular of winter buds, ofHydrangea macrophylla was examined. The non-flower-bearing shoot is usually composed of a lower and an upper part, between which a boundary is discernible by means of a distinctly short internode. This internode is the lowermost of the upper part, and it is usually shorter than the internodes immediately above and below, although the internodes tend to shorten successively from the proximal to the distal part of the shoot. Variations exist in the following characters among the terminal bud, the axillary bud on the lower part of the shoot and the axillary bud on the upper part: (1) length of bud; (2) character of the outermost pair of leaf primordia; (3) degree of development of secondary buds in the winter bud; and (4) the number of leaf primordia. Usually, the terminal bud contains several pairs of foliage leaf primordia with a primordial inflorescence at the terminal of the bud, but the axiallary bud contains only the primordia of foliage leaves in addition to a pair of bud scales.     

Branching pattern and inflorescence bud displacement in Flickingeria (Orchidaceae)

The vegetative architecture of Flickingeria is modular, consisting of three kinds of shoots of determinate growth. Species differ with respect to extent of branching and length of shoots, but branching is normally restricted to a few particular buds, the potential of which depends on their position.
All inflorescences are axillary, although some appear to be terminal. Inflorescences subtended by foliage leaves are displaced into a cavity in the stem, emerging on the abaxial side of the leaf base. The peripheral layers of the stem covering the inflorescence bud may be conspicuously extended and dry up so as to resemble a bract. The arrangement of the inflorescences is a distinctive specific character within the genus. A hypothesis for the evolution of character states is established. The growth and flowering strategy is discussed.     

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.     

Localization of axillary meristems during different stages of radish ontogenesis

An analysis of axillary meristem (axillary bud) localization of radish (Raphanus sativus L. cv. Tetra-I?ówiecka) was undertaken on vernalized (flowering) and unvernalized (vegetative) plants. It has been shown that the localization of these meristems can be different on successive nodes of the same plant and is connected with the development stages of the plants. The axillary meristems can arise on the stem as well as in the leaf axil or on the base of the subtending leaf. The localization of axillary meristems has been discussed in relation to growth directions and growth correlations inside the meristematic region of the shoot apex.     

中国水仙花芽分化观察及储藏条件对花芽数的影响研究

以三年生中国水仙‘金盏银台’为材料,采用石蜡切片法观察其花芽形态分化过程。结果表明：中国水仙的花芽分化从7月上旬开始,到9月中旬形成雌蕊结束。其过程可分为叶芽时期、花序原基形成期、佛焰状总苞形成期、花原基形成期、花冠形成期、雄蕊形成期、雌蕊形成期7个时期。其中花冠形成期较长,20 d左右。花芽的外部形态变化上,分化后期芽的生长速度明显快于前期。对鳞茎球内花序数量的统计结果显示,高温储藏及烟熏法共同使用对中国水仙花序的形成具有很好的促进作用。     

Reproduction site preference and performance by sexuparae,and mating behavior of their sexual generation on the primary host plant in a heteroecious aphid,Neothoracaphis yanonis (Homoptera: Aphididae)

We studied reproduction site preference and performance by the sexuparae (autumnal migrants) of Neothoracaphis yanonis and mating behavior of their sexual generation on its primary host plant, Distylium racemosum. The sexuparae preferred younger leaves of D. racemosum for settlement and imbibing leaf sap, and they produced more offspring there than on older leaves. Thus, it is suggested that the sexuparae selected more nutritious younger leaves to increase their own fecundity. The offspring consist of yellowish dark-grey and creamy yellow type nymphs, which develop into small males and large oviparae, respectively. Yellowish dark-grey nymphs were deposited gregariously on the basal part of the abaxial surface of a leaf blade, while creamy yellow nymphs were deposited evenly over the abaxial surface. Such a localized distribution pattern probably resulted from sexuparae's strategy to enhance a possibility for their male offspring to mate with oviparae at the base of leaf blade, over which oviparae crawl to twigs. During mating and sometimes afterwards, the ovipara incidentally carried a male on her back and crawled downward on twig to find an axillary bud for oviposition. This behavior may be advantageous to males, who can guard their mate during this period against other conspecific males. Such mate-guarding behavior seems to be related to the development of dual mate-seeking strategies, in which males try to copulate, first at the basal part of midrib, and second on the axillary bud.     

Anatomy of shoots and tumors of in vitro habituated Rhododendron 'Montego' (Ericaceae) cultures with tissue proliferation

Tissue proliferation (TP) is characterized primarily by the formation of galls or tumors at the crown of container-grown rhododendrons that were initially propagated in vitro. In the cultivar 'Montego', TP-like symptoms are first observed in vitro as shoot clusters with small leaves and nodal tumors. In addition, unlike the normal in vitro non-TP (TP-) shoots, in vitro TP (TP+) shoots proliferate rapidly without the presence of the plant growth regulator cytokinin in the tissue culture medium. Comparisons of the anatomy of TP+ and TP- shoot tips showed that TP+ shoots had a less developed vascular system, longer cells in the pith and cortex, and altered internodal elongation at the shoot apex. In addition, TP+ axillary buds were abnormal in that they were displaced onto the stem above the leaf axil, and a small group of proliferating cells replaced the shell zone at the base of the bud. Initiation of tumor formation began with the expansion of this region of cell proliferation (RCP) and shoot growth from the abnormal axillary bud (tumor bud). Organization of the tumor bud and extension of the RCP characterized the further development of two types of tumors. In polar shoot tumors, shoot growth continued from the persistent tumor bud and the tumor at the base of the shoot remained small in size. The RCP extends downward to the vascular junction of the subtending leaf and the stem of the TP+ shoot. In nonpolar tumors, continuous de novo meristem formation led to the development of large tumors with or without shoots. The RCP is present throughout the tumor and is associated with de novo meristem formation. Comparisons to the anatomy of other tumor-like structures showed that TP tumors of Rhododendron 'Montego' are most similar to tobacco genetic tumors.     

The inflorescence structure of Campylotropis (Leguminosae)

Racemose inflorescences were investigated organographically and ontogenetically in three species of Campylotropis. An additional phyllome was found to be borne at the base of the pedicel in C. hirtella and C. giraldii but not in C. polyantha. Organographic observations revealed that the additional phyllome is the prophyll on the reduced lateral branch borne in the axil of the phyllome on the central inflorescence axis, and that the additional phyllome subtends the pedicel terminating in a flower. The flower presents its abaxial side to the additional phyllome in accordance with the organographic observations. Ontogenetic observations, moreover, revealed the presence of a rudimentary apex of the reduced branch in earlier stages of development. On the basis of such evidence, the central inflorescence axis of Campylotropis is interpreted as being composed of reduced lateral branches each of which usually bears only one flower in the additional phyllome axil. The inflorescence structure is kept in spite of the loss of the additional phyllome. In comparison with allied genera, the inflorescence of Campylotropis is regarded as a reduced pseudoraceme. An evolutionary trend in the inflorescence structure of the genus is inferred.     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

DEVELOPMENT OF THE EPIPHYLLOUS INFLORESCENCE OF HELWINGIA JAPONICA (HELWINGIACEAE)

The inflorescence of Helwingia japonica (Thunb.) Dietr. is initiated adjacent to the leaf axil on the adaxial side of the base of a leaf primordium during its second plastochron. The inflorescence which develops from the resulting primordium comes to be situated on the midrib of the mature fertile leaf, through the action of a basal, intercalary meristem. In fertile leaves this meristem develops beneath, as well as above, the insertion of the inflorescence primordium on the leaf primordium. The same meristem is present in sterile leaves as well. A separate, adaxial vascular bundle departs from the leaf trace in the base of the petiole and leads to the inflorescence, in the mature fertile leaf. This adaxial vascular bundle is absent in sterile leaves. It is argued that the vascular anatomy does not conclusively confirm the hypothesis that the epiphyllous inflorescence is the congenital fusion product of a leaf and an axillary inflorescence. Instead, it is suggested that the interplay of changes in the position of primordium initiation, and intercalary growth, offers an ontogenetic explanation of the situation, which in turn may be related to the phylogeny of the species in question. It appears to be misguided and futile to look for homologies (i.e., 1:1 correspondences) between fertile and sterile leaves, since 1:1 correspondences do not exist in this case.