The development of pistillate and perfect florets in Xeranthemum squarrosum (Asteraceae)

The formation of capitulum inflorescence with two different types of floret is an interesting issue in floral biology and evolution. Here we studied the inflorescence, floral ontogeny and development of the everlasting herb, Xeranthemum squarrosum, using epi‐illumination microscopy. The small vegetative apex enlarged and produced involucral bracts with helical phyllotaxy, which subtended floret primordia in the innermost whorl. Initiation of floret primordia was followed by an acropetal sequence, except for pistillate peripheral florets. The origin of receptacular bracts was unusual, as they derived from the floral primordia rather than the receptacular surface. The order of whorl initiation in both disc and pistillate flowers included corolla, androecium and finally calyx, together with the gynoecium. The inception of sepals and stamens occurred in unidirectional order starting from the abaxial side, whereas petals incepted unidirectionally from the adaxial or abaxial side. Substantial differences were observed in flower structure and the development between pistillate and perfect florets. Pistillate florets presented a zygomorphic floral primordium, tetramerous corolla and androecium and two sepal lobes. In these florets, two sepal lobes and four stamen primordia stopped growing, and the ovary developed neither an ovule nor a typical stigma. The results suggest that peripheral pistillate florets in X. squarrosum, which has a bilabiate corolla, could be considered as an intermediate state between ancestral bilabiate florets and the derived ray florets.     

Floral Development in Sonja White Clover (Trifolium repens) a Papilionoid Legume

Floral development in Sonja white clover was examined usingscanning electron microscopy. Florets and bracts were foundto arise from common primordia initiated as protuberances fromthe apical meristematic area of the inflorescence. The patternof floret initiation on the inflorescence was acropetal, theoldest florets resting basally. Floral organ initiation withineach floret was acropetal, petals being initiated before stamens.Floret development was zygomorphic, each whorl of floral organsdeveloping unidirectionally from the abaxial side. There wasfound to be overlapping in the timing of initiation and developmentof these organs. Antesepalous stamens were found initially tooutgrow their antepetalous counterparts. Early petal developmentwas synpetalous. Eglandular hairs were found basally on thecalyx cup and on the pedicel. Procumbent hairs were found tobe more numerous and randomly distributed on the abaxial surfacesof the mature calyx cup. Trifolium repens L., Sonja cultivar, white clover, scanning electron microscopy, floral development, inflorescence     

Comparative ontogeny of perfect and pistillate florets in Senecio vernalis (Asteraceae)

We studied the inflorescence, and in particular ontogeny and development of the florets in Senecio vernalis as a representative member of Asteraceae, using epi-illumination microscopy. Initiation and subsequent development of florets on the highly convex inflorescence apex occur acropetally, except for pistillate ray florets, which show a lag in initiation. Receptacular bracts derive from the receptacular surface after development of all florets. The order of whorl initiation in both disc and ray florets include corolla, androecium and finally the pappus, together with the gynoecium. Development of corolla lobes from a ring meristem occurs in bidirectional order starting from the lateral side, whereas stamens incept unidirectionally from the abaxial side. Concurrently with the inception of two median carpel primordia, a ring meristem develops at the base of the corolla from which pappus bristles differentiate in later stages. Pistillate ray florets show significant differences from perfect disc florets as reflected by the zygomorphic shape of the floral apex and a shift of floral merosity from pentamery to tetramery. Loss of stamens in ray florets occurs due to abortion of primordia after initiation.     

REGENERATION OF THE SUNFLOWER CAPITULUM AFTER CYLINDRICAL WOUNDING OF THE RECEPTACLE

Using the young capitulum of Helianthus annuus L., a cylindrical plug of undifferentiated receptacle tissue, 1 mm in diameter, was isolated from lateral communication with the rest of the receptacle surface by a vertical circular wound cut, while retaining continuity with the subapical meristem. Within 24 hr, active cell division was induced at the inner and outer surfaces of the wound and in the receptacle epidermis bordering the wound edges, creating a rounded rim at the top of the wound. Within 3–6 days, floral initials, spaced 133–166 μm apart appeared on the flanks of both rims and later on the top of the plug and surrounding receptacle surface. The first formed initials developed into involucral bracts or ray florets and the later ones into disc florets which were organized into contact parastichies, the number of which did not conform with the Fibonacci series. The base of the plug developed into a stem-like structure completing the regeneration of a fully formed functional capitulum. This operation was demonstrated for two sunflower cultivars and occurred in both long and short daylengths.     

MISSING FLOWERS gene controls axillary meristems initiation in sunflower

The initiation and growth of axillary meristems are fundamental components of plant architecture. Here, we describe the mutant missing flowers (mf) of Helianthus annuus characterized by the lack of axillary shoots. Decapitation experiments and histological analysis indicate that this phenotype is the result of a defect in axillary meristem initiation. In addition to shoot branching, mutation affects floral differentiation. The indeterminate inflorescence of sunflower (capitulum) is formed of a large flat meristem which produces floret primordia in multiple spirals. In wildtype plants a bisecting crease divides each primordium in two distinct bumps that adopt different fate. The peripheral (abaxial) part of the primordium becomes a small leaf-like bract and the adaxial part becomes a flower. In the mf mutant, the formation of flowers at the axil of bracts is precluded. Histological analyses show that in floret primordia of the mutant a clear subdivision in dyads is not established. The primordia progressively bend inside and only large involucral floral bracts are developed. The results suggest that the MISSING FLOWERS gene is essential to provide or perceive an appropriate signal to the initiation of axillary meristems during both vegetative and reproductive phases.     

INFLORESCENCE AND FLORAL DEVELOPMENT IN HOUTTUYNIA CORDATA (SAURURACEAE)

The inflorescence of Houttuynia cordata produces 45–70 sessile bracteate flowers in acropetal succession. The inflorescence apical meristem has a mantle-core configuration and produces “common” or uncommitted primordia, each of which bifurcates to form a floral apex above, a bract primordium below. This pattern of organogenesis is similar to that in another saururaceous plant, Saururus cernuus. Exceptions to this unusual development, however, occur in H. cordata at the beginning of inflorescence activity when four to eight petaloid bract primordia are initiated before the initiation of floral apices in their axils. “Common” primordia also are lacking toward the cessation of inflorescence apical activity in H. cordata when primordia become bracts which may precede the initiation of an axillary floral apex. Many of these last-formed bracts are sterile. The inflorescence terminates with maturation of the meristem as an apical residuum. No terminal flowers or terminal gynoecia were found, although subterminal gynoecia or flowers in subterminal position may overtop the actual apex and obscure it. Individual flowers have a tricarpellate syncarpous gynoecium and three stamens adnate to the carpels; petals and sepals are lacking. The order of succession of organs is: two lateral stamens, median stamen, two lateral carpels, median carpel. The three carpel primordia almost immediately are elevated as part of a gynoecial ring by zonal growth of the receptacle below the attachment of the carpels. The same growth elevates the stamen bases so that they appear adnate to the carpels. The trimerous condition in Houttuynia is the result of paired or solitary initiations rather than trimerous whorls. Symmetry is bilateral and zygomorphic rather than radial. No evidence of spiral arrangement in the flower was found.     

The Effect of Population Density on Floret Initiation, Development and Abortion in Winter Wheat

There is little information in the literature concerning thephysiological basis of the relationship between plant populationdensity and kernel number in winter wheat (Triticum aestivumL.). Thus, two experiments were conducted to evaluate this relationship.Expt 1, involving three population densities, was carried outnear Taian, China in 1982 and in Expt 2, two densities wereevaluated near Lexington, Kentucky in 1986. Plants were sampled every 2 d in the spring, main stem spikeswere dissected and florets were scored according to a 10-stagescale of development. The rate of primordia initiation increasedas density increased until the point at which primordia numberswere equal in all treatments. After this point, an increasein density reduced the primordia initiation rate. In both experimentsincreasing density reduced the total number of floret primordiainitiated and the number of kernels per spike. In Expt 1 theeffect of density on kernel number per spike was accounted forapproximately equally by the effect of density on number ofprimordia initiated and floret abortion. In Expt 2, however,floret abortion was influenced much less by density and accountedfor only 7 % of the variation in kernel number per spike. Thekey result was that the effect of density was determined earlyin floral development. The data suggest that yield losses athigh densities may be determined too early in development tobe offset by N applications at the terminal spikelet stage. Triticum aestivum L., spike development, spikelet development, seeding rate     

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.     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

ORGAN INITIATION AND DEVELOPMENT OF INFLORESCENCES AND FLOWERS OF ACACIA BAILEYANA

Inflorescence and floral ontogeny are described in the mimosoid Acacia baileyana F. Muell., using scanning electron microscopy and light microscopy. The panicle includes first-order and second-order inflorescences. The first-order inflorescence meristem produces first-order bracts in acropetal order; these bracts each subtend a second-order inflorescence meristem, commonly called a head. Each second-order inflorescence meristem initiates an acropetally sequential series of second-order bracts. After all bracts are formed, their subtended floral meristems are initiated synchronously. The sepals and petals of the radially symmetrical flowers are arranged in alternating pentamerous whorls. There are 30–40 stamens and a unicarpellate gynoecium. In most flowers, the sepals are initiated helically, with the first-formed sepal varying in position. Petal primordia are initiated simultaneously, alternate to the sepals. Three to five individual stamen primordia are initiated in each of five altemipetalous sectorial clusters. Additional stamen primordia are initiated between adjacent clusters, followed by other stamens initiated basipetally as well as centripetally. The apical configuration shifts from a tunica-corpus cellular arrangement before organogenesis to a mantle-core arrangement at sepal initiation. All floral organs are initiated by periclinal divisions of the subsurface mantle cells. The receptacle expands radially by numerous anticlinal divisions in the mantle at the summit, concurrently with proliferation of stamen primordia. The carpel primordium develops in terminal position by conversion of the floral apex.     

FLORAL DEVELOPMENT IN GYMNOTHECA CHINENSIS (SAURURACEAE)

The development of the inflorescence and flowers are described for Gymnotheca chinensis Decaisne (Saururaceae), which is native only to southeast China. The inflorescence is a short terminal spike of about 50–70 flowers, each subtended by a small bract. There are no showy involucral bracts. The bracts are initiated before the flowers, in acropetal order. Flowers tend to be initiated in whorls of three which alternate with the previous whorl members. No perianth is present. The flower contains six stamens, and four carpels fused in an inferior ovary containing 40–60 ovules on four parietal placentae. Floral symmetry is dorsiventral from inception and throughout organ initiation. Floral organs are initiated in the following order: 1) median adaxial stamen, 2) a pair of lateral common primordia which bifurcate radially to produce two stamen primordia each, 3) median abaxial stamen, 4) a pair of lateral carpel primordia, 5) median adaxial carpel, 6) median abaxial carpel. This order of initiation differs from that of any other Saururaceae previously investigated. The inferior ovary results from intercalary growth below the level of stamen attachment; the style elongates by intercalary growth, and the four stigmas remain free. The floral structure of Gymnotheca is relatively advanced compared to Saururus, but its assemblage of specializations differs from that of either Anemopsis or Houttuynia, the other derived genera in the Saururaceae.     

Inflorescence and flower development in the Hedychieae (Zingiberaceae): Hedychium coccineum Smith

The inflorescence of Hedychium coccineum Smith is thyrse, and the primary bracts are initiated in a spiral phyllotactic pattern on the sides of the inflorescence dome. Cincinnus primordia are initiated on the flank of the inflorescence apex, in the axils of primary bracts. This primordium subsequently develops a bract and a floral primordium. Then, the floral primordium enlarges, flattens apically, and becomes rounded. Sepals are initiated sequentially from the rounded corner of the primordium ring sepal initiation, and the floral primordium continues to enlarge and produces a ring primordium. Later, this ring primordium separates three common primordia surrounding a central cavity. The adaxial common primordium is the first separation. This primordium produces the posterior petal and the fertile stamen. The remaining two common primordia separate and produce respectively a petal and a petaloid, the inner androecial member. As the flower enlarges, the cavity of the floral cup becomes a rounded–triangular apex; these apices are the sites of outer androecial primordium initiation. The abaxial outer androecial member slightly forms before the two adaxial members develop. But this primordium ceases growth soon after initiation, while the two posterior primordia continue growth to produce the lateral petaloid staminodes. During this stage, gynoecial initiates in the floral cup and continues to grow until extending beyond the labellum.     

Arabidopsis thaliana (L.) Heynh. as a model object for studying genetic control of morphogenesis

A vast amount of information on the genetic control of plant development has been obtained in Arabidopsis thaliana with classical genetic and molecular biological methods. The genes involved in multistep regulation of floral morphogenesis have been identified. The formation of floral meristem is controlled by the LEAFY (LFY), UNUSUAL FLORAL ORGANS (UFO), APETALA1 (AP1), and APETALA2 (AP2) genes. Studies of the abruptus and bractea recessive monogenic mutants from the collection of the Department of Genetics and Selection, Moscow State University, showed that the ABRUPTUS (ABR) and BRACTEA (BRA) genes also play an important role in inflorescence differentiation. The ABR gene controls the early formation of organ primordia on the inflorescence and the formation of floral organ primordia after floral initiation. Further differentiation of inflorescence organ primordia in vegetative or generative organs depends on the activity of the LFY gene, and floral organ identity is determined by the homeotic genes. Presumably, the major function of the ABR gene is to determine the auxin polar transport. The BRA gene suppresses the development of bracts on the inflorescence and constrains cell division at the base of primordia of rosette and cauline leaves.     

Inflorescence and floral development in Astragalus lagopoides Lam. (Leguminosae: Papilionoideae: Galegeae)

Inflorescence and floral organogenesis and development of the bushy perennial legume Astragalus lagopoides of the section Hymenostegis were studied by means of epi-illumination light microscopy. Based on our observations, the primordia of lanceolate racemose inflorescences are born in the axils of leaves. Each inflorescence apex initiates acropetally bracts and floral apices for some time and then eventually ceases meristematic activity and forms an oblong-shaped terminal structure. The formation of such atypical terminal protrusion on the inflorescence meristem is judged to be a diagnostic feature for well-organized cessation of meristem morphogenesis. Pentamerous perfect flowers of the plant show strong zygomorphy and marked overlap in time of initiation among different organ primordia. Unexpectedly, sepal initiation is bidirectional starting from the lateral sides of the floral apex. Other significant developmental feature includes the existence of two types of common primordia, which are formed successively. From the primary common primordia there are produced antesepalous stamens and secondary common primordia. In comparison, the five secondary common primordia subdivide into a petal and an antepetalous stamen primordia. Initiation of two different types of common primordia is possibly the result of rising overlap in time of initiation of organs and demonstrates an advanced developmental style in the genus Astragalus.     

Heterochronic development of the floret meristem determines grain number per spikelet in diploid, tetraploid and hexaploid wheats

Background and Aims

The inflorescence of grass species such as wheat, rice and maize consists of a unique reproductive structure called the spikelet, which is comprised of one, a few, or several florets (individual flowers). When reproductive growth is initiated, the inflorescence meristem differentiates a spikelet meristem as a lateral branch; the spikelet meristem then produces a floret meristem as a lateral branch. Interestingly, in wheat, the number of fertile florets per spikelet is associated with ploidy level: one or two florets in diploid, two or three in tetraploid, and more than three in hexaploid wheats. The objective of this study was to identify the mechanisms that regulate the architecture of the inflorescence in wheat and its relationship to ploidy level.

Methods

The floral anatomy of diploid (Triticum monococcum), tetraploid (T. turgidum ssp. durum) and hexaploid (T. aestivum) wheat species were investigated by light and scanning electron microscopy to describe floret development and to clarify the timing of the initiation of the floret primordia. In situ hybridization analysis using Wknox1, a wheat knotted1 orthologue, was performed to determine the patterning of meristem formation in the inflorescence.

Key Results

The recessive natural mutation of tetraploid (T. turgidum ssp. turgidum) wheat, branching head (bh), which produces branched inflorescences, was used to demonstrate the utility of Wknox1 as a molecular marker for meristematic tissue. Then an analysis of Wknox1 expression was performed in diploid, tetraploid and hexaploid wheats and heterochronic development of the floret meristems was found among these wheat species.

Conclusions

It is shown that the difference in the number of floret primordia in diploid, tetraploid and hexaploid wheats is caused by the heterochronic initiation of floret meristem development from the spikelet meristem.Key words: Triticum, wheat, inflorescence, spikelet, floret, meristem, heterochrony, heterochronic development, knotted1, polyploidy     

CONTROL OF INTERCALARY GROWTH IN THE SCAPE OF GERBERA BY AUXIN AND GIBBERELLIC ACID

With the inflorescence removed, intercalary growth can be maintained in the scape of Gerbera jamesonii by application of gibberellic acid (GA, gibberellin A₃) or indole-3-acetic acid (IAA); the latter usually promotes more rapid and greater elongation than the former because of a greater effect on older tissues. Simultaneous application of the two substances, even when both are at optimal levels, promotes more rapid elongation than either substance alone; in fact, the rate of elongation may equal that of the intact scape. In decapitated scapes (receptacle and involucral bracts removed with the inflorescence), GA and IAA promote cell elongation with reduced or no cell division. In deflowered scapes (receptacle and involucral bracts intact) both GA and IAA promote cell division, as well as cell elongation, so that the pattern of scape elongation is nearly the same as that for intact scapes. Apparently the bracts and receptacle contribute something required for cell division which acts in concert with GA and IAA. Deflowered and decapitated scapes elongate at nearly the same rates initially; thus the rate of elongation does not depend on cell division. The ultimate length of the scape is dependent on cell number and, hence, cell division, since deflowered scapes attain greater lengths than those that are decapitated.     

鹅耳枥和虎榛子（桦木科）雌性生殖器官的形态发生

为了进一步理解类群之间的系统发育关系,在扫描电镜下,对桦木科植物鹅耳枥（Carpinus turczaninowii Hance)和虎榛子（Ostryopsis davidiana Decne.)的雌花序、小花序和雌花的原基形成和发育过程进行了观察。两种植物均具单性花、雌雄同株。其雌花序为复合的穗状花序,每两朵小花构成一个小聚伞花序,多个这样的小花序螺旋排列在一个总花序轴上。小花序由5枚苞片组成,1枚初级苞片,4枚次级苞片。后分别由两个半环状的共同原基发育而来。鹅耳枥的近轴面次级苞片生物缓慢,远轴面的生长较快,成熟时呈扩展的叶片状;虎榛子的近轴面和远轴面次给苞片均生长较快,成熟时靠全呈囊状。花被原基为环状,花被随着子房的发育而而逐渐长大包围子房并与之愈合。研究对前人有关小花序的苞片数目和两个二心皮子房的定位方式等方面的认识作了澄清或纠正。每个小花序的苞片数目不像Abbe观察的那么多;二心皮子房定位方式在鹅耳枥属是互成直角,而非相互平行。     

Morphogenesis of Female Reproductive Organs in Carpinus turczaninowii and Ostryopsis davidiana (Betulaceae)

For better understanding of the relationships between genera, the primordium occurrence and morphological developmental process of female inflorescence, cymule and floret in Carpinus turczaninowii Hance and Ostryopsis davidiana Decne. of the Betulaceae were observed under the scanning electron microscope (SEM). Both species were monoecious. Their female inflorescence was a compound spike comprising several cymules arranged helically along an inflorescence axis. Each cymule consisted of two florets and five bracts, i.e., one primary bract and four other secondary ones which were developed from two semi-circular common primordia, respectively. In Carpinus , the adaxial secondary bracts grew slowly, while the abaxial ones grew fast, resulting in the appearance of a wide leafy bract upon maturity. In Ostryopsis , however, both abaxial and adaxial secondary bracts were fully developed, becoming a bladder-like but unclosed involucre when mature. Perianth primordia in both genera were circular. When the ovary became larger and larger, the perigone grew gradually, and finally surrounded and was adnate to the ovary. Some traditional viewpoints on the number of bracts and the orientation of bicarpellate ovary in cymule were clarified based on this study. The cymule bracts were not so many as those observed by Abbe; and the two bicarpellate ovaries were orientated perpendicularly, rather than parallel.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

HETEROMORPHIC FLOWER DEVELOPMENT IN NEPTUNIA PUBESCENS,A MIMOSOID LEGUME

The characteristic of heteromorphic inflorescences in some mimosoid legumes such as Neptunia is a puzzling one which can be approached developmentally. Each spicate inflorescence of Neptunia pubescens includes three types of flowers: perfect in the upper half, functionally male just below the middle, and sterile or neuter at the base. Developmental studies of the inflorescence show that order of initiation of bracts on the inflorescence is acropetal, but that order of subsequent development of flowers is both acropetal and basipetal on the axis. Bract growth and initiation of the axillary floral apices at the base are inhibited or retarded, while those in the middle and upper levels continue development without interruption. The three types of floral primordia are similar during initiatory stages of organ formation and through early development. At mid-development, differences arise in floral symmetry, petal form, stamen form, and size and shape of the carpel. The functionally male flowers become strongly dorsiventral and zygomorphic while the other two morphs remain actinomorphic or nearly so. Heteromorphy arises from a combination of early suppression of organogeny plus mid-stage innovations of zygomorphy and lateral expansion of stamen primordia. These divergent developmental pathways in one inflorescence can be interpreted in part using Gould's concept of heterochrony: changes in timing of developmental events to produce different structures. Other changes in Neptunia cannot be explained by this concept, however; such changes as omission of processes (i.e., meiosis) in some organs, or addition of processes not normally present (i.e., blade formation in stamen primordia which become staminodia). It is becoming evident from work on this and other legume flowers that actual loss of organs is rare, compared to initiation followed by suppression or modification.