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.     

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 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.     

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.     

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.     

INFLORESCENCE AND FLOWER DEVELOPMENT IN THE PIPERACEAE. II. INFLORESCENCE DEVELOPMENT OF PIPER

The inflorescence development of three species of Piper (P. aduncum, P. amalago, and P. marginatum), representing Sections Artanthe and Ottonia, was studied. The spicate inflorescences contain hundreds or even thousands of flowers, depending on the species. Each flower has a tricarpellate syncarpous gynoecium and 4 to 6 free stamens, in the species studied. No sepals or petals are present. In P. marginatum the apical meristem of the inflorescence is zonate in configuration and is unusually elongate: up to 1,170 μm high and up to 480 μm wide during the most active period of organogenesis. Toward the time of apical cessation both height and diameter gradually diminish, leaving an apical residuum which may become an attenuate spine or may be cut off by an abscission zone just below the meristem. The active apex produces bract primordia; when each is 40–55 μm high, a floral apex is initiated in its axil. Both bract and floral apex are initiated by periclinal divisions in cells of the subsurface layer. The bracts undergo differentiation rather early, while the floral apices are still developing. The last-produced bracts near the tip of the inflorescence tend to be sterile.     

毛舞花姜花器官的发生与发育

通过扫描电镜观察了毛舞花姜(Globba barthei Gagne p.)的花序及花器官的发生与发育。3枚萼片原基首先于花顶连续发生,随后花顶的中心凹陷形成环状原基,环状原基进一步分化形成三枚花瓣—雄蕊共同原基,并在花顶的中心形成花杯。共同原基分化形成花瓣和三枚内轮雄蕊,紧接着外轮雄蕊在花杯的顶点发生。远轴的两枚内轮雄蕊延伸生长并相互融合形成了唇瓣,近轴的一枚形成了可育雄蕊;近轴的两枚外轮雄蕊发育形成了成熟花结构中的侧生退化雄蕊,而远轴的一枚缺失。近轴的两枚外轮雄蕊原基起始的同时,3枚心皮原基也在中心花杯的内侧发生而后与外轮雄蕊相间排列。对毛舞花姜花序的发生和发育的观察发现,在花序轴的头几片初级苞片中产生的是珠芽原基而非蝎尾状小花序原基,其形态特征类似于早期的蝎尾状小花序原基,由此推测珠芽很可能是蝎尾状小花序的变异。     

Flower development inAntirrhinum majus L. (Scrophulariaceae) with a comment upon corolla tube formation

The inflorescence apex is stratified and has a single layered tunica, bracts as well as all the floral organs develop acropetally. Except for stamens which are initiated with the activity of deeper layers of apex, all other floral appendages are initiated with the activity of second layer. On the basis of ontogenetic studies the floral organs have been homologised with the leaves. The adjacent calyx lobes develop and remain closely appressed and do not show post-genital fusion. Corolla tube is formed due to zonal growth preceeded by joint growth of hump on the abaxial face of stamen primordia and interprimordial region between petal primordia. Thus no post-genital fusion was observed. The placentae develop concomitantly with carpellary wall.     

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.     

Inflorescence and Flower Development of Globba barthei ( Zingiberaceae)

Inflorescence of Globba barthei is a thyrse . Primary bracts are initiated in a spiral phyllotactic pattern on the inflorescence apex . Cincinnus primordia are initiated in the axils of primary bracts . These promordia develop secondarybracts and floral primordia . The floral primordium continues to enlarge and produce a ring primordium . Sepals are initiated sequentially from the rounded corner of the primordium . The ring primordium separates three common primordium surrounding a central cavity . The adaxial common primordium is the first to separate . This primordium divides transversely and producespetal and fertile stamen . The remaining two common primordium transversely separate and produce respectively a petal and a petaloid . As the flower developing , the cavity of the floral cup becomes triangular . The angles of this triangle are the sites of outer androecial primordium . The abaxial androecia forms slightly earlier than the two adaxial ones, and then this primordium ceases growth soon . The two posterior primordia continue growth to produce the lateral petaloid staminodes . During this stage , gynoecia initiate from the floral cup and continue to fuse and develop into style and stigma. In addition ,Initiation of the bulbil primordium is observed at base of inflorescence axis during the early floral development . The bulbil primordium initiates in the axil of primary bract . The evolutionary significance of six androecia is discussed .     

Floral and inflorescence morphology and ontogeny in Beta vulgaris, with special emphasis on the ovary position

Background and Aims

In spite of recent phylogenetic analyses for the Chenopodiaceae–Amaranthaceae complex, some morphological characters are not unambiguously interpreted, which raises homology questions. Therefore, ontogenetic investigations, emphasizing on ‘bracteoles’ in Atripliceae and flowers in Chenopodioideae, were conducted. This first paper presents original ontogenetic observations in Beta vulgaris, which was chosen as a reference species for further comparative investigation because of its unclarified phylogenetic position and its flowers with a (semi-)inferior ovary, whereas all other Chenopodiaceae–Amaranthaceae have hypogynous flowers.

Methods

Inflorescences and flowers were examined using scanning electron microscopy and light microscopy.

Key Results

Floral development starts from an inflorescence unit primordium subtended by a lateral bract. This primordium develops into a determinate axis on which two opposite lateral flowers originate, each subtended by a bracteole. On a flower primordium, first five tepal primordia appear, followed by five opposite stamen primordia. Simultaneously, a convex floral apex appears, which differentiates into an annular ovary primordium with three stigma primordia, surrounding a central, single ovule. A floral tube, which raises the outer floral whorls, envelops the ovary, resulting in a semi-inferior ovary at mature stage. Similarly, a stamen tube is formed, raising the insertion points of the stamens, and forming a staminal ring, which does not contain stomata. During floral development, the calyces of the terminal flower and of one of the lateral flowers often fuse, forming a compound fruit structure.

Conclusions

In Beta vulgaris, the inflorescence is compound, consisting of an indeterminate main axis with many elementary dichasia as inflorescence units, of which the terminal flower and one lateral flower fuse at a later stage. Floral parts develop starting from the outer whorl towards the gynoecium. Because of the formation of an epigynous hypanthium, the ovary becomes semi-inferior in the course of floral development.Key words: Beta vulgaris, Chenopodiaceae, floral ontogeny, gynoecial development, epigynous hypanthium, semi-inferior ovary, inflorescence ontogeny, LM, SEM     

Three new species of Mitrephora (Annonaceae) from Sabah, Malaysia

Three new species of Mitrephora (Blume) Hook. f. & Thomson, M. clemensiorum, M. vittata and M. woodii , are described from Sabah, Malaysia. Mitrephora clemensiorum is related to M. korthalsiana Miq., but is distinguished by having long inflorescences, long flowering and fruiting pedicels, and fruits with sessile monocarps. Mitrephora vittata is related to M. reflexa Merr., but has lanceolate, subcoriaceous leaves, and densely pubescent inflorescence rachides, flowering pedicels and bracts. Mitrephora woodii is similar to M. heyneana (Hook. f. & Thomson) Thwaites, but has smaller flowers and fruits with sessile monocarps     

ONTOGENY OF THE INFLORESCENCE OF SAURURUS CERNUUS (SAURURACEAE)

The spicate inflorescence of Saururus cernuus L. (Saururaceae) results from the activity of an inflorescence apical meristem which produces 200–300 primordia in acropetal succession. The inflorescence apex arises by conversion of the terminal vegetative apex. During transition the apical meristem increases greatly in height and width and changes its cellular configuration from one of tunica-corpus to one of mantle (with two tunica layers) and core. Primordia are initiated by periclinal divisions in the subsurface layer. These are “common” primordia, each of which subsequently divides to produce a floral apex above and a bract primordium below. The bract later elongates so that the flower appears borne on the bract. All common primordia are formed by the time the inflorescence is about 4.4 mm long; the apical meristem ceases activity at this stage. As cessation approaches, cell divisions become rare in the apical meristem, and height and width of the meristem above the primordia diminish, as primordia continue to be initiated on the flanks. Cell differentiation proceeds acropetally into the apical meristem and reaches the summital tunica layers last of all. Solitary bracts are initiated just before apical cessation, but no imperfect or ebracteate flowers are produced in Saururus. The final event of meristem activity is hair formation by individual cells of the tunica at the summit, a feature not previously reported for apical meristems.     

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.     

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.     

THE ONTOGENY OF THE INFLORESCENCE AND FLOWER OF LIQUIDAMBAR STYRACIFLUA L. (HAMAMELIDACEAE)

A developmental study of the inflorescence of Liquidambar styraciflua L. was conducted to clarify morphological discrepancies reported in the literature. Salient features of development are: 1) the inflorescence apex results from the conversion of a terminal, vegetative apex; 2) partial inflorescence apices arise as ellipsoid structures in axils of leaves, bracts, or transitional phyllomes; 3) development of male heads is acropetal whereas female heads differentiate basipetally; 4) the partial inflorescence apex becomes segmented into several distinct subunits indicating an axillary branch system of the third order; 5) distinct individual floral primordia are initiated on the subunits; 6) a complete absence of perianth development; 7) inception of carpel primordia in flowers of lower male heads as well as female heads, but a failure of the gynoecium to develop beyond an incipient stage in male heads; and 8) development of sterile structures around the base of the styles of only female flowers near the time of anthesis. Carpellary characteristics of the sterile structures are described, their morphological nature is discussed, and the phylogenetic position of Liquidambar is evaluated.     

大戟科麻疯树属三种植物花器官发生

利用扫描电子显微镜观察了大戟科Euphorbiaceae麻疯树属Jatropha麻疯树J. curcas L.、佛肚树J. podagrica Hook.和棉叶麻疯树J. gossypifolia L.花器官发生。结果表明: 麻疯树、佛肚树和棉叶麻疯树花萼原基均为2/5型螺旋发生。在同一个种不同的花蕾中, 花萼的发生有两种顺序: 逆时针方向和顺时针方向。远轴面非正中位的1枚先发生。5枚花瓣原基几乎同时发生。雄花中雄蕊两轮, 外轮对瓣, 内轮对萼。研究的3种麻疯树属植物雄蕊发生方式有两种类型: 麻疯树亚属麻疯树的5枚外轮雄蕊先同时发生, 5枚内轮雄蕊后同时发生, 佛肚树亚属佛肚树和棉叶麻疯树雄蕊8-9枚, 排成两轮, 内外轮雄蕊同时发生。雌花的3枚心皮原基为同时发生。麻疯树属单性花, 雌花的子房膨大而雄蕊退化, 雄花的雄蕊正常发育, 子房缺失。根据雄蕊发生方式, 支持将麻疯树属分为麻疯树亚属subgen. Jatropha和佛肚树亚属subgen. Curcas。     

INFLORESCENCE AND FLOWER DEVELOPMENT IN THE PIPERACEAE. I. PEPEROMIA

Flowers of Peperomia species are the simplest structurally of any of the members of the Piperaceae. The spicate inflorescences form terminally and in axillary position; in each, the apex first is zonate in configuration with a two-layered tunica while 3-4 leaves are initiated. Later, when the inflorescence apical meristem begins bract initiation, the biseriate tunica persists, but zonal distinctions diminish and the apex can be described in terms of a simple tunicacorpus configuration. The inflorescence apex aborts after producing 30-40 bracts in acropetal succession an abscission layer forms across the base of the apex, and the meristem dries and drops off. Bracts are produced by periclinal divisions in T₂ (and occasionally also in the third layer as well); the later-formed floral apices arise by periclinal divisions in T₂ and the third layer. Each floral apex is at first a long transverse ridge in the axil, perpendicular to the long axis of the inflorescence. This establishes bilateral symmetry in the flower, which persists throughout subsequent growth. The floral meristem becomes saddle-shaped, and two stamen primordia are delimited, one at either end and lower than the central floral apex. A solitary carpel is initiated abaxially, and soon forms a circular rim which heightens as a tube with an apical pore. Within the open carpel, a solitary ovule is initiated from the entire remains of the floral apical meristem; it, hence, is terminal in the flower, and its placentation is basal. Carpellary closure in P. metallica results from accelerated growth of the abaxial lip, and the two margins become appressed. Species differ greatly as to whether the abaxial or the adaxial lobe predominates in late stages of carpel development. In P. metallica, the receptive portion of the stigma forms from the shorter lobe which is overtopped. Stigmatoid tissue forms internal to the receptive stigma. The prevailing bilateral floral symmetry, absence of a perianth, and the spicate inflorescence are features which distinguish Peperomia (and Piperaceae) from the magnolialian line of angiosperms.     

Wound-Induced Initiation of Involucral Bracts and Florets in the Developing Sunflower Inflorescence

The production of additional floral organs by the inflorescenceof Helianthus annuus as a boron deficiency symptom was examinedand found to be related to the occurrence of minute splits inthe receptacle of the young capitulum. Wounding the receptacleby puncturing or cutting at an early stage when the receptacledome was forming (floral stage 3) or later when the receptaclehad either become saucer-shaped (floral stage 4) or the firstrows of disc floret primordia were appearing on the rim of thereceptacle (floral stage 5) resulted in the initiation of involucralbracts, ray and disc florets in the wound area, reproducingthe symptoms of boron deficiency. When the receptacle was woundedat later floral stages, when the receptacle was partially orwholly covered by disc floret primordia, involucral bracts andray florets were not formed in the proximity of the wound, leadingto the conclusion that the commitment of floral organ primordiabegins at the time of their initiation. The inductive effectsof wounding could not be reproduced by indoleacetic acid, naphthaleneaceticacid or benzyladenine applied to the receptacle surface in lanolinepaste. The results are related to the normal development ofthe sunflower inflorescence and it is concluded that the firstfloral organ primordia that appear in floral stage 3 and developinto involucral bracts during floral stage 4 may determine thepositions of subsequent primordia and establish the radial symmetryof the inflorescence. ¹ Supported by a grant from the Australian Research Grants Scheme. (Received August 10, 1982; Accepted October 12, 1982)     

马桑绣球(绣球科)的花器官发生和发育

在扫描电镜下观察了马桑绣球Hydrangea aspera孕性花的发生及发育过程。马桑绣球的花器官向心轮状发生：花萼原基以2/5螺旋式相继发生,花瓣原基几乎同步发生。花瓣开始发育时,与花萼相对的雄蕊发生。与花瓣相对的雄蕊原基与心皮原基几乎同时出现。初始心皮向上扩展,分化出花柱和柱头,向下延伸,嵌入花托,发育为下位子房。花发育成熟时,隔膜于子房的下部连续,而中部和上部不连续,即子房为不完全2室。经过与绣球属已观察过的另外5种1亚种花器官发生和发育比较,发现马桑绣球与藤绣球H. ano mala subs