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.     

PHYLLODE THEORY IN RELATION TO LEAF ONTOGENY IN SANSEVIERIA TRIFASCIATA

Shoot apices of Sansevieria trifasciata have a three-layered mantle, a zone of subapical initials, a central meristem, and a peripheral meristem. Leaf initiation begins with periclinal divisions in L-3 and is followed by periclinal divisions in L-2 and anticlinal divisions in L-l. At first, the primordium is a mound of tissue at one point on the flank, but it soon takes the form of a low ridge encircling the apex. An ephemeral adaxial meristem differentiates in L-2 of the primordium when it is about 50 μ high and is active until the primordium is about 450 μ high. Then it ceases basipetally and is not observable after the primordium is about 600μ high. As the adaxial meristem ceases at the base of the radial tip, its two lateral regions become the submarginal meristems of the expanded portion. Marginal meristems differentiate from the protoderm, and oblique-anticlinal divisions of the marginal initials result in the formation of an abaxial and adaxial epidermis. These derivatives undergo a few anticlinal divisions, increasing marginal width, and then they divide periclinally, increasing marginal thickness. After the primordium is about 600-700 μ high it continues to grow in length by a diffuse basal intercalary meristem. When the leaf is 3 dm long, an adaxial rounding meristem differentiates in the region just above the sheath. Leaf vasculature consists of parallel bundles which anastomose acropetally. Vascular bundles are arranged in a semicircle in the expanded portion and in a circle in the radial tip. There is one centrally located bundle at the apex as a result of lateral anastomoses. Present evidence from leaf ontogeny and mature vasculature in S. trifasciata is interpreted as supporting the concept that the liliaceous leaf is homologous with the phyllodes of A corns and Acacia.     

木立芦荟叶的发育解剖学研究

应用植物解剖学方法研究了木立芦荟（Aloe arborescens Mill.)叶的发育过程。研究结果表明,叶原基在发育早期其形态是不对称的,内部为同形细胞组成,但很快分化成原表皮,原形成层束和基本分生组织。以后,原表皮发育成表皮,位于原表皮下的2－5层基本分生组织细胞发民同化薄壁组织,而位于中央的基本分生组织细胞则发育成储水薄壁组织,原形成层束发育成维管束。维管束由维管束鞘、木质部、韧皮部和大型薄壁细胞组成。大型薄壁细胞起源于原形成层束,位于韧皮部内,其发育迟于筛管、伴胞,为芦荟属植物叶的结构特征。     

EARLY HISTOGENESIS OF THE ADULT LEAVES OF DARLINGTONIA CALIFORNICA (SARRACENIACEAE) AND ITS BEARING ON THE NATURE OF EPIASCIDIATE FOLIAR APPENDAGES

In order to assess the validity of various interpretations of tubular leaves of angiosperms, a histogenetic study of the ontogeny of adult leaves of Darlingtonia californica was undertaken. The adult leaf of Darlingtonia is characterized by a sheathing leaf base, an elongate ascidium, an overarching hood, and two “fishtail” appendages which arise near the leaf apex. A keellike growth, with two rows of alternate vascular bundles, traverses the tube from base to mouth. Ontogenetic studies show that the primordium arises by a monopodial rather than a sympodial mode of growth as previously reported. After the formation of a small, erect primordium, a restricted adaxial meristem is initiated that expands both adaxially and upwards. This “querzone” serves, in effect, to congenitally combine the two primordial margins during its extension. Growth and maturation of the subjacent portions cause tubular elongation in the leaf. Primordial apical divisions are later replaced by more general intercalary growth with acropetal and centrifugal maturation. The hood and fishtails are established early in ontogeny by adaxial growth of the primordial apex and subsequent activation of juxtaposed localized meristems. Comparative morphology has established that the epiascidiate leaf is a foliar appendage that undergoes certain specific morphogenetic modifications. It has a structural relationship to ensiform appendages of Acacia and Acorus as well as to peltate foliar organs. The early ontogeny of Darlingtonia leaves is considered to be homologous with other epiascidiate foliar organs, including some supposedly primitive carpels.     

Untersuchungen über den Ursprung der fertilen Fieder vonOphioglossum pedunculosum

Observations of young leaf primordia give information about the origin of the fertile spike ofOphioglossum pedunculosum Desv. Each primordium shows a certain asymmetry that is visible in form and position of the fertile spike primordium, but above all in the course of the marginal meristem. The fertile spike primordium is connected with the marginal meristem on the right or that on the left side of the sterile segment. On the basis ot these observations the following concept of the origin of the fertile spike is formed: The marginal meristem curves on one side of the leaf primordium and turns towards the ventral side. This process is followed by meristem fractionation in the course of which the smaller part curved towards the middle of the leaf primordium becomes independent and initiates the development of the fertile spike, whereas the larger part of the marginal meristem contributes to the growth of the sterile segment.

     

LEAF DEVELOPMENT IN DOXANTHA UNGUIS-CATI (BIGNONIACEAE)

Leaf structure in Doxantha unguis-cati is polymorphic. The usual mature compound leaf is composed of two lanceolate leaflets and a terminal tripartite spine-tendril. Leaf primordia are initiated simultaneously in pairs on opposite flanks of the shoot apical meristem by periclinal cell divisions in the third subsurface layer of the peripheral flank meristem. Two leaflet primordia are the first lateral appendages of the compound leaf. Initiation of these leaflet primordia occurs on the adaxial side of a compound leaf primordium 63–70 μm long. Lamina formation is initiated at the base of a leaflet primordium 70–90 μm long and continues acropetally. Mesophyll differentiation occurs in later stages of development of leaflets. The second pair of lateral appendages of the leaf primordium differentiate as prongs of the tendril. Initiation of the second pair of lateral appendages occurs on the adaxial side of a primordium approximately 168 μm long. Acropetal procambialization and vacuolation of cells extend to the apex of tendrils about 112 μm long, restricting the tendril meristem to the adaxial side of the primordium and resulting in curvature of the tendril. The tendril meristem is gradually limited to a more basipetal position as elongation of apical cells continues. Initiatory divisions and early ontogenetic stages of leaflets and tendrils are similar. Their ontogeny differs when the lateral primordia are approximately 70 μm long. Marginal and submarginal initials differentiate within leaflets but not in tendrils. Apical growth of tendrils ceases very early in ontogeny as compared with leaflets.     

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

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

The Structure of the Plumule of Nelumbo Nucifera Gaertn. and the Nature of its Scale

The structure of the plumule of Nelumbo nucifera Gaertn. and its feature covered with scale are seldom seen in dicotyledon. The fact that the plumule possesses scale is even more uncommon. This particular phenomenon is investigated by observing the differentiation of the plumule apex and the development of the leaf organs. After the seed is formed, the embryo has two young leaves and a terminal bud covered with scale. In the bud it has already differentiated the 3rd and the 4th leaf primordium and a shoot apex, the differentiation of which is very complex. So the structure of the plumule passes through 4 plastochrons altogether. It is made clear through observation and analysis that, before the 4th leaf primordium is formed, the transforma- tions of the shoot apex of the embryo in each plastochron are fundamentally alike. After the 4th leaf primordium is developed, the shoot apex becomes complex and there appear 3 different active cell regions which become the bases of vegetative bud of the seeding apex. The development of these 3 active cell regions will be stated in “The Structure of the Vegetative Bud of Nelumbo nucifera Gaertn. and the Nature of its Scales.” The apices of the plumule are almost slightly domed in structure. As a rule, their width is from 95 to 107 μ. Their height is from 17 to 20 μ during one plastochron. Before the 3rd leaf initiation, the anatomical structure of apices is examined and the fol- lowing zones may be delimited: zone of tunica initials, zone of corpus initials, peripheral zone, and zone of rib meristems. It is frequently observed that the cell of corpus in subapical peripheral zone develops periclinal division, which is the initial cell of leaf primordium; Procambium will appear before the stage of the appearance of leaf buttress. The apex of the plumule is in an apical position, but when the seedling is formed, as the developing leaves are alternate, the directions of the shoot apex are changed, simultaneously the base part of the leaf encloses the axis, and the adaxial meristem also differentiates the scale which encloses the terminal bud, thus placing the bud in axillary of the leaf and forming a zigzag phenomenon of the axis of the seedling. Above the basal adaxial side of the leaf primordium develops the scale of the plumule with meristem periclinal division of closely attached protoderm as its base. So the scale of the plumule of Nelumbo nucifera Gaertn. and the axillary stipule are of the same origin. To sum up, the scale of the embryo of Nelumbo nucifera Gaertn. is differentiated from the adaxial meristem of the basal part of the leaf primordium, and is the derivative part of the leaf. It has the same function as the coleoptile of the monocotyledon. Whether they are homologous organs or not is still to be investigated.     

Intrapetiolar inflorescence buds in Salacca (Palmae): development and significance

The inflorescence in all species of Salacca is enclosed in a chamber within the leaf base and is exserted through a slit on the abaxial surface of the leaf base. The inflorescence bud is interpreted ds an axillary meristem that becomes radially displaced by adaxial growth of the leaf primordium. A fine channel is produced from the leaf axil to the base of the inflorescence and persists at maturity. The channel and the bud chamber enlarge as the leaf elongates. They are lined by an epidermal layer. There is no cellular breakdown until the collapse and tearing of tissues of the leaf during inflorescence enlargement late in ontogeny. The vegetative bud is positioned about 130⁰ from the axil of its subtending leaf and lies directly below the abaxial inflorescence slit of the leaf above. Vegetative bud development was not observed, hut there is a suggestion of relatively late initiation. The separation of. Eleiodoxa from Salacca is supported by differences in the development of inflorescence and vegetative buds.     

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.     

ANATOMY AND ASPECTS OF DEVELOPMENT OF THE STAMINATE INFLORESCENCES AND FLORETS OF SEVEN SPECIES OF ALLOCASUARINA (CASUARINACEAE)

The morphology, ontogeny, and vascular anatomy of the staminate inflorescences and florets of seven species of Allocasuarina are described. The generally terminal but open-ended inflorescences occur on monoecious or staminate dioecious trees and consist of whorls of bracts, each subtending a sessile axillary floret. Each floret consists of one terminal stamen with a bilobed, tetrasporangiate anther enclosed typically by cuculliform appendages, commonly considered bracteoles, an inner median pair and an outer lateral pair. The mature stamen is exerted, the anther is basifixed and is extrorsely dehiscent. In early development of a male inflorescence very little internodal elongation occurs and enclosing cataphylls appear. The inflorescence apex is a low dome with a uniseriate tunica and a small group of central corpus cells. Bract primordia are initiated by periclinal divisions of C₁ followed by further divisions of the corpus and anticlinal divisions in the tunica. The bracts are epinastic and become gamophyllous except apically by cell divisions in both sides of each primordium. Stomata are restricted to the axis furrows and the abaxial tips of the bracts. The axillary florets arise in acropetal succession initiated by periclinal divisions in C₁ accompanied by anticlinal divisions in the tunica. The lateral floral appendages are also initiated by C₁ followed by anticlinal divisions in the tunica. They become adnate basally later with the subtending bract. The median sterile appendages are initiated in a manner similar to the initiation of the outer appendages. The stamen is initiated by divisions in the outer layers of the corpus and in the tunica, and then develops first by apical growth followed by intercalary growth. The vascular system of the inflorescence is identical to that of the vegetative stem. Each floret is supplied by a single bundle that has its source in a branch from each of the two traces supplying a bract. Six bundles arise from the floral bundle; four of these terminate in the base of the stamen and two form an amphicribal bundle that supplies the anther. Pollen is binucleate, 3- to 7-porate. The exine is tegillate.     

STRUCTURE AND DEVELOPMENT OF LEAVES IN CARLUDOVICA PALMATA (CYCLANTHACEAE) WITH REFERENCE TO OTHER CYCLANTHACEAE AND PALMAE

The adult leaf of Carludovica palmata consists of a plicate lamina, adaxial hastula, petiole, and sheath. The leaf is unusual in the angiosperms because about two-thirds of the apical meristem is utilized in its initiation. The adult leaf requires about 4–5 plastochrons to mature. Shortly after its initiation the adult leaf and apical meristem collectively appear pyramid-shaped and various parts of the mature adult leaf may be traced back to particular portions of the pyramid. Plications develop by differential growth within the lamina, not by splitting of leaf tissue. Quantitative studies indicate that certain regions of the developing adult leaf elongate more rapidly or slowly than other regions depending upon the stage of leaf development. The adult leaf of C. palmata develops differently from those of previously studied palms in various ways. It therefore appears less justifiable to consider the superficial similarity between the adult leaves of various Cyclanthaceae (particularly those of Carludovica sensu strictu) and those of fan palms as evidence of especial affinity between the Cyclanthaceae and Palmae. Juvenile leaves of C. palmata differ from adult leaves both in their mode of origin and appearance at maturity. The juvenile leaf appears homologous to the entire adult leaf.     

Developmental study onHypolepis punctata (Thunb.) Mett.

The origins of the first and second petiolar buds ofHypolepis punctata were clarified in relation to the early development of the leaf primordium, which arises from a group of superficial cells of the shoot apical meristem. One of these superficial cells produces a two-sided leaf apical cell which subsequently cuts off segments to make a well-defined cell group, called here the leaf apical cell complex, on the distal part of the leaf primordium. Meanwhile, cells surrounding the leaf apical cell complex also divide frequently to form the basal part of the leaf primordium. Two groups of basal cells of the leaf primordium located on the abaxial and the adaxial sides initiate the first and the second petiolar buds, respectively. The initial cells are usually contiguous to the leaf apical cell complex, constructing the abaxial and adaxial flanks of the very young leaf primordium. However, the first petiolar bud sometimes develops from cells located farther from the leaf apical cell complex. These cells are derived from those originally situated in the peripheral region of the shoot apical meristem. This study was supported by a Grant-in-Aid for Encouragement of Young Scientists by the Ministry of Education, Science and Culture, of Japan No. 474322 in 1979.     

LEAF DEVELOPMENT IN SENECIO ROWLEYANUS (COMPOSITAE)

Prolonged apical growth of the leaf primordium and the presence of distinct marginal meristems do not occur in Senecio rowleyanus. Intercalary cell divisions accompanied by radial expansion of derivatives from an adaxial meristem account for the spherical shape of the leaf. The “window” in the lamina marks the position of the adaxial meristem and precludes interpretation of the leaf as being unifacial. Stomata are mesoperigenous and anomocytic in type. Schizogenous secretory canals occur in both the leaf and the stem, and their association with vascular bundles is discussed. The anatomy of the leaf is interpreted in terms of xeromorphy.     

Initiation of axillary and floral meristems in Arabidopsis

Shoot development is reiterative: shoot apical meristems (SAMs) give rise to branches made of repeating leaf and stem units with new SAMs in turn formed in the axils of the leaves. Thus, new axes of growth are established on preexisting axes. Here we describe the formation of axillary meristems and floral meristems in Arabidopsis by monitoring the expression of the SHOOT MERISTEMLESS and AINTEGUMENTA genes. Expression of these genes is associated with SAMs and organ primordia, respectively. Four stages of axillary meristem development and previously undefined substages of floral meristem development are described. We find parallels between the development of axillary meristems and the development of floral meristems. Although Arabidopsis flowers develop in the apparent absence of a subtending leaf, the expression patterns of AINTEGUMENTA and SHOOT MERISTEMLESS RNAs during flower development suggest the presence of a highly reduced, "cryptic" leaf subtending the flower in Arabidopsis. We hypothesize that the STM-negative region that develops on the flanks of the inflorescence meristem is a bract primordium and that the floral meristem proper develops in the "axil" of this bract primordium. The bract primordium, although initially specified, becomes repressed in its growth.     

ONTOGENY AND PHYLLOTAXIS OF THE TERMINAL VEGETATIVE SHOOTS OF MICHELIA FUSCATA

Tucker Shirley C. (Northwestern U., Evanston, Ill.) Ontogeny and phyllotaxis of the terminal vegetative shoots of Michelia fuscata. Amer. Jour. Bot. 49(7): 722–737. Illus. 1962.—Two patterns of symmetry occur in Michelia fuscata In the lead shoots, leaves arise in a 2/5 spiral arrangement which may be either clockwise or counterclockwise. Other shoots are dorsiventrally organized; these shoots produce leaves in a modified ½ phyllotaxis in which the angle between the 2 files of leaves lies between 100° and 150°, according to the particular branch. Both types of shoot have a zonate apical meristem with a biseriate tunica a central initial zone, and a peripheral zone. The apical configuration of cells does not change appreciably during the plastochron. The flat to low-convex outline of the shoot apex is maintained by initiation of the leaves close to the summit of the apex; the diameter of the meristem diminishes greatly after such an initiation. Leaf inception in the subsurface tunica layer is followed by precocious activity of the marginal meristems which extend the stipular flanges completely around the base of the apical meristem. The stipular margins then fuse laterally and form a hood over the apex. A subapical initial meanwhile is active in the leaf blade, where it persists up to the time the leaf is 2 mm high. The most recent primordium is 300 μ high before another leaf is initiated. The vascular system of the stem is a cylindrical network of leaf traces, with 6–12 traces per leaf. The procambium develops acropetally from preexisting vascular strands in the stem below. Elements of the diverse sclereid system differ in shape in different tissues, according to the availability of intercellular space. Goebel's term “Pendelsymmetrie” is discussed with reference to apical activity in Michelia.     

榛属（桦木科）花序及花的形态发生

在扫描电镜下观察了桦木科榛属榛、毛榛和滇榛的花序和花的形态发生过程。榛属雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基;每个花原基分化出2个心皮原基,形成二心皮雌蕊;雌蕊基部有2层花被原基,内层花被原基环状,外层花被发生于花原基近轴面和远轴面,近轴面和远轴面的花被不均等分化,外层花被发生早于内层花被。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出2枚次级苞片和4。6个雄蕊原基,形成4—6枚雄蕊,每个雄蕊具4个药囊,在雄蕊原基分化形成4药囊雄蕊过程中．出现雄蕊原基纵裂。并且花丝纵裂至基部。为进一步全面探讨桦木科属间系统演化关系提供了证据。     

榛属 (桦木科) 花序及花的形态发生

在扫描电镜下观察了桦木科榛属榛、毛榛和滇榛的花序和花的形态发生过程。榛属雌花序由多个小聚伞花序螺旋状排列组成;每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化形成2个花原基;每个花原基分化出2个心皮原基,形成二心皮雌蕊;雌蕊基部有2层花被原基,内层花被原基环状,外层花被发生于花原基近轴面和远轴面,近轴面和远轴面的花被不均等分化,外层花被发生早于内层花被。雄花序为柔荑状,由多个小聚伞花序螺旋状排列组成。每个小花序原基分化出1枚初级苞片和一团小花序原基分生组织,由小花序原基分生组织分化出2枚次级苞片和4~6个雄蕊原基,形成4~6枚雄蕊,每个雄蕊具4个药囊,在雄蕊原基分化形成4药囊雄蕊过程中,出现雄蕊原基纵裂,并且花丝纵裂至基部。为进一步全面探讨桦木科属间系统演化关系提供了证据。     

THE ANATOMY OF THE PISTILLATE INFLORESCENCE AND FLOWER OF CASUARINA VERTICILLATA LAMARCK (CASUARINACEAE)

The pistillate inflorescence of Casuarina verticillata is described as consisting of a primary axis bearing whorls of bracts with a cymule in the axil of each bract of the more central whorls. Each cymule consists of an atepallate, two-carpellate, syncarpous floret and two, lateral, once-lobed bracteoles. A “peripheral intercalary” meristem, in which divisions are primarily periclinal, forms a meshwork beneath the bracts from early development and moves the connate bracts centrifugally around the cymules and extends and binds the bracts, and to some extent the bracteoles, of the fertile part of the inflorescence together. Each bract receives a single trace; each cymule receives two traces. Each bundle extension of a cymule trace supplies: 1) a branch which joins its counterpart to become the anterior common carpellary bundle; 2) a second branch which joins its counterpart to become the posterior common carpellary bundle; and 3) a central branch which supplies a lateral bracteole. Within each floret, each common carpellary bundle provides a dorsal carpellary bundle, two ventral carpellary bundles (fertile anterior carpel) or one common ventral bundle (sterile posterior carpel). The ventral bundle-supplies join and form a single placental bundle which lies in the gynoecial septum, and which, in turn, supplies the two ovules in the anterior carpel. Whether the inflorescence is a simple racemose or a condensed cymose type cannot be determined from this species alone. The function of the sclerenchymatous, enclosing bracteoles and connate bracts is discussed.     

Model for the role of auxin polar transport in patterning of the leaf adaxial–abaxial axis

Leaf adaxial–abaxial polarity refers to the two leaf faces, which have different types of cells performing distinct biological functions. In 1951, Ian Sussex reported that when an incipient leaf primordium was surgically isolated by an incision across the vegetative shoot apical meristem (SAM), a radialized structure without an adaxial domain would form. This led to the proposal that a signal, now called the Sussex signal, is transported from the SAM to emerging primordia to direct leaf adaxial–abaxial patterning. It was recently proposed that instead of the Sussex signal, polar transport of the plant hormone auxin is critical in leaf polarity formation. However, how auxin polar transport functions in the process is unknown. Through live imaging, we established a profile of auxin polar transport in and around young leaf primordia. Here we show that auxin polar transport in lateral regions of an incipient primordium forms auxin convergence points. We demonstrated that blocking auxin polar transport in the lateral regions of the incipient primordium by incisions abolished the auxin convergence points and caused abaxialized leaves to form. The lateral incisions also blocked the formation of leaf middle domain and margins and disrupted expression of the middle domain/margin‐associated marker gene WUSCHEL‐RELATED HOMEOBOX 1 (SlWOX1). Based on these results we propose that the auxin convergence points are required for the formation of leaf middle domain and margins, and the functional middle domain and margins ensure leaf adaxial–abaxial polarity. How middle domain and margins function in the process is discussed.